ys aoe Walls a oat ed vas Death vi phitts . Feige at ih Aan Sia, diay eth a oi oe ‘ ayia ihe As at Hy herred fad Ave oH eh “ been fated hy saat te ates ae. ‘i Mak sbs tal “ af ait oe ; eh tts fh fa . * yeaa ie p Ha ri a fs heal , LT UA wt abe arty) ee Se eh eee sh at ay ere satan 9b ah Aanak etn Gena igrenye (teeta at rh has dye ICE, he 4 a dee ede k Bt a { wang Bre te fa Mtoe Da vallnl gh a : see i i ita le iad ihe in oat, ef Mada ta hao dA 1 64 ahh bh YS aN be rele vi ae ha th irs gea trang Pt eee Siateeate ip Mere hep pad kt Aa heath AG 4 q < oH 1 Vn SAS aN) 0h wy re! Bact cr 9e 1 Koh ORES: * ah Z ‘ai eared Ps abt a oe a SPs q YM ah ia tat seh ete oes + A SECM ADEE % oh 28 bab oS 1440 Maley tp enter pisces oe pod 2 SF LI2AT TM ERIE: ray pee M03 exe ras ece?: )\ we AFATSON! yey sa Journal of the Botanical Society of the British Isles Volume 26 Part 1 February 2006 Editors: M. Briggs, M.J. Y. Foley, D. L. Kelly, D. R. McKean, M. S. Porter, C. D. Preston, M. N. Sanford Botanical Society of the British Isles Registered Charity Number: 212560 Applications for membership should be addressed to the Hon. General Secretary, c/o Department of Botany, The Natural History Museum, Cromwell Road, London, SW7 5BD, from whom copies of the Society’s Prospectus may be obtained. Officers for 2005-2006 President, Mr R. D. Pryce Vice-Presidents, Prof. C. A. Stace, Prof. A. J. Richards, Mr A. O. Chater, Dr C. D. Preston Acting Honorary General Secretary, Mr D. A. Pearman Honorary Treasurer, Mr M. E. Braithwaite Editors of Watsonia Papers and Notes, M. J. Y. Foley, D. L. Kelly, D. R. McKean, M. N. Sanford* Book Reviews, C. D. Preston Plant Records, M. S. Porter Obituaries, M. Briggs *Receiving editor, to whom all MSS should be sent (see inside back cover). © 2006 Botanical Society of the British Isles The Society takes no responsibility for ue views expressed by authors of Papers, Notes, Book Reviews or Obituaries. The cover illustration of Potentilla anglica Laich. (Trailing Tormentil) was drawn by Rosemary Wise. Watsonia 26: 1—7 (2006) Sorbus whiteana (Rosaceae), a new endemic tree from Britain TC. G, RICH* Department of Biodiversity and Systematic Biology, National Museums & Galleries of Wales, Cardiff CF 10 3NP and L. HOUSTON School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG ABSTRACT Sorbus whiteana T. C. G. Rich & L. Houston, a new endemic tree from the Avon Gorge and Wye Valley, is described. It is distinguished from other members of the S. aria (L.) Crantz group in Britain by the obovate, unlobed leaves which are greenish under- neath, and the fruits being longer than wide. It is triploid. About 76 plants are known in four populations and a plant from another locality require confirmation. The complex history of its discovery is set out; it has been confused with S. wilmottiana E. F. Warb. and S. hungarica (Bornm.) Hedl. ex C. E. Salmon. KEYWORDS: Avon Gorge, Bristol, Sorbus aria group, Whitebeam. INTRODUCTION The Avon Gorge in Bristol is home to at least nine native and four introduced taxa of Sorbus, Whitebeams, and is the richest and most important Sorbus site in Britain. Of the rarer species, S$. bristoliensis Wilm. and _ S. wilmottiana E. F. Warb. are endemic to the Avon Gorge, S$. eminens E. F. Warb and S. porrigentiformis E. F. Warb. are endemic to England and Wales, and S. anglica Hedl. is endemic to Britain and Ireland. The commoner native species in the Avon Gorge are S. aria (L.) Crantz sensu stricto, S. aucuparia L. and S. torminalis (L.) Crantz, and the introduced species are §S. croceocarpa P. D. Sell, S. decipiens (Bechst.) Irmisch, S. intermedia (Ehrh.) Pers. and S. latifolia (Lam.) Pers. In this paper we describe a ninth native Sorbus from the Avon Gorge (the fifth species there that is endemic to Britain). It is a member *e-mail: tim.rich@nmgw.ac.uk of the S. aria group, of which four other members also occur in the Avon Gorge (S. aria sensu stricto, S. eminens, S. porrigentiformis and S. wilmottiana). It has been known from the Avon Gorge for some time but not differentiated; it has also recently been found in the Wye Valley. METHODS Broad leaves from the short vegetative shoots, excluding the oldest and youngest leaf (Aas et al. 1994) were measured on herbarium material in NMW. Fruits were measured on fresh material, and the colours matched against the Royal Horticultural Society colour charts (Royal Horticultural Society 1966). Potential pollen viability was investigated using Alexander’s Stain (Alexander 1969) on the three flowering collections available. Anthers were removed from herbarium specimens with tweezers under a low-power binocular microscope, and placed on a slide with a drop of Alexander’s Stain, warmed briefly on the hotplate, then broken up with the tweezers. The preparation was then covered with a coverslip and replaced on the hotplate to improve the uptake of the stain. The slides were then examined under a_ high-power compound microscope for areas of dense pollen grains. Potentially viable grains were counted as those which were large and rounded- triangular with cell walls which stained green and with cytoplasm inside which stained uniformly bright red. Small deformed grains or those staining green only with very little or no red staining inside (i.e. no cytoplasm) were considered infertile. 2) T. C. G. RICH & L. HOUSTON Chromosome numbers were determined using standard methods. Tips of actively growing roots were collected mid-morning and pre-treated in 8-hydroxylquinoline solution for 20-24 hours at 4°C. They were then fixed in a 3:1 mixture of absolute ethanol: glacial acetic acid and stored at 4°C until needed. Root tips were then hydrolysed in concentrated 5N HCl for 10 minutes at room temperature, then stored in 70% industrial methylated spirits. Meristems were dissected out in 45% acetic acid, and squashed between a microscope slide and coverslip in 0 1% aceto-orcein stain with brief flame treatment. The distribution was assessed from herbaria (BIRM, BM, BRISTM, CGE, K, NMW, OXF and UPS; abbreviations following Kent & Allen 1984), and from extensive Sorbus field work in South-west England and South Wales between 1996 and 2005. RESULTS A small amount of morphological variation was found within and between populations. Some plants from the Avon Gorge have relatively large leaves, and the plant from Lady Park Wood had relatively more veins. Typical leaves from the short shoots are illustrated in Figure 1. Potential pollen fertilities were found of 43% for one plant on the top northern edge of Great Quarry, Clifton Down (voucher NMW. V.2002.17.189), 68% for another plant in woodland nearby (NMW.V.2004.3.109), and 62% for a plant from the Great Quarry (NMW. V.2003.1.154, = Lemche’s aria33), giving an overall mean of 58%. A few giant pollen grains (cf. Liljefors 1955) were noted in the latter collection. Chromosome counts of 2n = 51 (triploid) have been obtained for a single plant grown from seed collected from the holotype tree in Quarry 3 in the Avon Gorge, and from Ban-y- gor cliffs (voucher NMW.V.2002.17.103) (J. Bailey, pers. comm. 2004). The potential pollen fertility is perhaps higher than might be expected for a triploid. Sorbus whiteana T. C. G. Rich & L. Houston, sp. nov. HOLOTYPE: Near top centre of Quarry 3, Leigh Woods, Avon Gorge, Bristol, v.c. 6 North Somerset, 21 August 2002, T. C. G. Rich & L. Houston (NMW, accession number V.2002.17.50). ISOTYPE: BM. Vernacular name: White’s Whitebeam. Frutex vel arbor parva ad 10 m ut minimum. Petioli 9-20 mm. Folia lata brachyblastorum (8 5—)9 3-13 6(-148) x (50)-5 6-8 7(-9 6) cm, (1 3—-)1 4-1 9(-20) plo longiora quam latiora, obovata vel obtrullata, ad longitudinis suae (53—)55-65(-69) partes centensimas latissima, apice acuto vel obtuso, basi cuneata, plerumque non lobata vel interdum (praecipue in umbra) leniter lobata, marginibus uniserratis vel leniter biserratis, dentes prope apicem extra sed in parte inferiore magis prorsum projectos ferentibus, sed c. 1 cm infimo prope petiolum fere integro; pagina superior glabrescens viridis, pagina inferior cinereoviridis tomentosa, venarum paribus (7 5—)8 5-11 5(- 125) ad angulum (24—)26—35(—37)° a costa tentis. Inflorescentia aggregata, tholiformis, ramulis viridialbis tomentosis instructa. Flores ad 17 mm lati. Sepala anguste triangularia, patentia, tomentosa sed ad apicem viridia, eglandulata. Petala (6—)7—-10 x 5-7 mm, cir- cularia vel late elliptica ungue brevi, alba, cupulata, pilis effusis ad basin vestita. Stamina c. 20, antheris luteis e roseo suffusis. Styli 2, virides, usque ad basin discreti, basi pilosi. Fructus maximi (9 5—)10—13 5(-14) x (9-)10—- 13(-14) mm, (0 85-)0 9-1 1(-1 2) plo longiores quam latiores, plerumque aspectu longiore quam latiore, interdum globosi, in medio latissimi, lenticellis sparsis vel satis frequentibus parvis vel mediocribus super superficiem dispersis, interdum ad basin aliquot pluribus, in maturitate rubri. Shrub or small tree to at least 10 m. Petioles 9-20 mm. Broad-leaves of short shoots (8 5—) 9 3-13 6(-148) x (5 0-)5 6-8 7(-96) cm, (1 3-)1 4-1 9(-2 0) times as long as wide, obovate to obtrullate, widest at (53—)55—65(— 69)% along leaf length, with apex acute to obtuse and base cuneate, usually unlobed or sometimes (especially in shade) weakly lobed, with margins uniserrate to weakly biserrate, with teeth at the top directed outwards, those below directed more forwards, but with lowest c. | cm near petiole nearly entire; upper surface glabrescent, green, lower surface greyish-green tomentose, with (7 5—)8 5-11 5(—12 5) pairs of veins held at an angle of (24—)26—35(—37)° to midrib. Inflorescence crowded, domed, with branchlets greenish-white tomentose. Flowers up to 17 mm across. Sepals narrowly tri- angular, spreading, tomentose but green at tip, eglandular. Petals (6—)7—10 x 5—7 mm, circular to broadly elliptic with a short claw, white, SORBUS WHITEANA 3 A } B D E F G H FIGURE 1. Sorbus whiteana. A-H, Typical leaves from the short shoots. A, Leigh Woods, Holotype and B-C, Isotype. D, Stokeleigh Camp (NMW.V.2004.26.183). E, Clifton Down, top north edge of Great Quarry (NMW.V.2001.25.240). F, Ban-y-gor (NMW.V.2002.17.103). G, Shorn Cliff (NMW.V.2002.13.164). H, Lady Park Wood (V.2004.11.36). I, Fruits, The Gulley (NMW.V.1998.48.21). Scale bars 1 cm. 4 T. C. G. RICH & L. HOUSTON py N (he) Ross-on-Wye Wales 2 ey ) a’ 3 Monmouth CS (Ks Ge. se) < Chepstow a | V.c. 34 West Gloucester ”A England ve} ot sf Bristol: 722 FIGURE 2. Distribution map of Sorbus whiteana in South-west England and Wales. @ Confirmed localities, © Provisional identification. SORBUS WHITEANA 5 cupped, with spreading hairs at base. Stamens c. 20, anthers yellow with a pink flush. Styles 2, green, free to base, hairy at base. Largest fruits (9-5—)10-13-5(-14) x (9-)10-13(-14) mm, (0-85—)0-9-1-1(-1-2) times as long as wide, usually looking longer than wide, sometimes globose, widest at the middle, with sparse to fairly frequent, small to moderate- sized lenticels scattered over the surface, sometimes with a few more at the base, red at maturity (RHS guide 44A, 45B, 46A, 46C). Sorbus whiteana occurs on Carboniferous Limestone rocks and their associated soils on open cliffs, screes and grassland edges, scrub and developing woodland. It is confined to the Avon Gorge, Bristol and the Wye Valley (Figure 2). All the NMW material cited below has been collected by either one or both of us, sometimes with other botanists and colleagues. V.C. 6 NORTH SOMERSET Avon Gorge, Leigh Woods, in three places: One tree, cliffs below Stokeleigh Camp, ST5673, 2004 (NMW), second tree nearby, 2005 (NMW); two trees, Donkey Slide, ST5673, 2004 (NMW); one tree, Quarry 3 (former police firing range), ST5574, 2002, 2004 (Holotype; NWVW). V.C. 34 WEST GLOUCESTER Avon Gorge, Clifton Down: Rocky wooded Slope, 1920, J. W. White (K). 13 September 1955, E. F. Warburg (OXF). 24 June 1958, P. J. M. Nethercott (OXF). We have seen it in three places: five trees, Great Quarry, ST5674, 2002, 2003, 2004 (NMW); at least 11 trees, wood north of Great Quarry ST5674, 1997, 2001, 2002, 2004 (NMW) and six more trees, 2005; trees 83 and 101x, north side of The Gulley, ST5674, 1996, 1997 (NMW; Rich & Houston 1996). A few trees, Ban-y-gor cliffs, Wye Valley, ST5496, 2000, 2002 (NMW). At least 53 plants, frequently scattered along Shorn Cliff, Tidenham, Wye Valley ST5398, ST5498 and ST5499, 2001, 2002 (NMW; Houston et al. 2002). V.C. 35 MONMOUTH One plant on cliffs, Lady Park Wood, Wye Valley, SO5414, 2003 (NMW;; Houston et al. 2004); requires confirmation from fruit. Thus at least 76 plants of S. whiteana are known in four sites (some with subpopulations) in two vice-counties in England, and it may occur in one site in Wales. DISCUSSION Sorbus whiteana is characterised by the obovate, cuneate-based, unlobed leaves which are greenish underneath and the red fruits which are generally longer than wide. Being triploid, S. whiteana is presumably derived from S. aria sensu stricto and another member of the S. aria group, either in the Wye Valley or in the Avon Gorge. Sorbus whiteana is only likely to be confused with the most closely related species in the S. aria group in the British Isles. The obovate, cuneate-based leaves which are greenish under- neath will distinguish it from most plants of the very variable S. aria sensu stricto which tends to have ovate leaves which are cordate to broadly cuneate at the base, and which are always white underneath. Sorbus whiteana differs from S. wilmottiana in having obovate, unlobed leaves which are greenish underneath; the latter has elliptic, weakly-lobed leaves which are whitish underneath (Rich & Houston 2004). The greenish underside of the leaves also distinguish S. whiteana from S. leptophylla E. F. Warb., which also has larger, shallowly- lobed leaves and larger fruits, and from S. vexans E. F. Warb., which also has relatively narrow leaves. Sorbus whiteana 1s readily distinguished from S. hibernica E. F. Warb., S. lancastriensis E. F. Warb., S. porrigentiformis and S. rupicola (Syme) Hedl. in having fruits longer than wide and leaves greener under- neath; these four other species have fruits wider than long and leaves whitish underneath. The complex history of S. whiteana in the Avon Gorge is worth setting out as the taxon has been known for some time. It was originally confused with the central European endemic S. hungarica (Bornm.) Hedl. ex C. E. Salmon in Journal of Botany 68: 174 (1930) (=S. hungarica (Bornm.) Karpati in Borbasia Nova 25: 23 (1944)) which is related to S. austriaca (G. Beck) Hedl. (Warburg & Karpati 1968). There has also been confusion with S. wilmottiana. Sorbus whiteana was first collected in the Avon Gorge by the great Bristol botanist James Walter White (1846- 1932), in 1920, from a rocky wooded slope on Clifton Down (K; there is no material in J. W. White’s herbarium in BRISTM), and our specific epithet commemorates him. The sheets were labelled ‘S. hungarica (Bornm.) Hedl.’ by C. E. Salmon, presumably after matching them with similar material named as such by T. 6 T.CsGFRICH Sale HOUSTON Hedlund from the Wye Valley (Salmon 1930; see also below). Other botanists attempted to collect White’s ‘S. hungarica’ but all the material traced so-labelled is S. aria, e.g. Mrs C. I. Sandwith & Mrs M. L. Wedgwood, 2 June 1931; Mrs C. I. Sandwith & G. Taylor, 20 September 1931 and perhaps more significantly two sheets from Clifton Down collected by E. F. Warburg no. 135, September 1935 (all BM) indicating that Warburg had tried to investigate it during his work on Sorbus. Warburg visited Clifton Down on several occasions and on 13 September 1955 collected three sheets of material, of which one is S$. whiteana and two others with narrower, more acute, more deeply lobed leaves are S. aria (OXF). These sheets were retained in a separate folder at OXF labelled ‘S. hungarica’ by Warburg, along with another sheet he had collected from Leigh Woods which may be part of the S. porrigentiformis group but which interestingly he had labelled “‘hungarica’ = ? wilmottiana’. Indeed it is probable that Warburg first discovered S. wilmottiana during his investi- gation of White’s “S$. hungarica’, but Warburg does not appear to have come to any firm conclusion about the *S. hungarica’ before he died in 1966. Warburg appears to have named at least one other S. whiteana collection (Clifton Down, 24° June 1958) P22 Jy 7M. Nethercott, OXF) as S. wilmottiana, but the specimen is mixed on a sheet with two genuine collections of S. wilmottiana. In 1996, English Nature commissioned a survey of Sorbus of The Gulley, Durdham Down, Bristol (v.c. 34 West Gloucester) to provide information before conservation work was carried out, as The Gulley was thought to be an important site for Sorbus (Rich & Houston 1996). During this survey, two plants of S. whiteana were found (another tree may be referable to the S. latifolia (Lam.) Pers. group) and were shown to P. D. Sell at Cambridge on 10 October 1996; he agreed they did not match any known species. The plants were given the working name Sorbus ‘JWWhite’ for convenience. Specimens were exhibited at the 1996 BSBI Exhibition Meeting (Rich & Houston 1997), which prompted only one comment, i.e. that they were S$. wilmottiana by P. J. M. Nethercott (this was not surprising in view of the determination by Warburg cited above). Between 1997 and 1999, L. Houston with others found more S. whiteana trees below the S. wilmottiana type tree on the edge of Clifton Down and collected further specimens, which were then all considered to be part of the variation in §$. wilmottiana (NMW). In 1996, Elin Lemche began a PhD on Sorbus taxonomy at Cambridge and, guided by L. Houston, collected Sorbus from the Avon Gorge for analysis using a variety of methods including DNA techniques (Lemche 1999). Amongst the specimens collected were two trees in secondary scrub from the bottom of the Great Quarry (Lemche nos. aria33 and aria34, locality incorrectly cited as Leigh Woods) which had at one time been demonstrated by various botanists as S. wilmottiana, although more recently they were known not to be that taxon. In her RFLP analysis, Lemche found that these two trees formed an isolated distinct group and in-situ hybridisation with Malus probes indicated that they were polyploid with one strong and one weak S. aria sensu lato band; she suggested that they were candidates for recognition as a new microspecies (Lemche 1999). However, P. D. Sell had named one as | S. rupicola (a species not currently known from the Avon Gorge) and the other was treated as S. aria. In 2001, T. C. G. Rich began to extend the work of Lemche (1999) on Sorbus taxonomy, using more advanced DNA methods (e.g. Fay et al. 2002), and requested to see the aria33 and aria34 trees that L. Houston had shown to E. Lemche. The trees were compared against the type tree of S. wilmottiana, and then against the 1996 Sorbus ‘JWWhite’ trees (ie. S. whiteana) and were found to be a good match for the latter. Further samples were collected from other trees in the area. It was then apparent that there had been long-standing confusion between S. whiteana and S. wilmottiana, at least on the Clifton side of the Gorge (see also Lemche 1999). Further detailed studies of Sorbus populations on the Leigh Woods side of the Avon Gorge (V.c. 6 North Somerset) by L. Houston in 2001 also clarified that the S$. whiteana plants did not fit within S. wilmottiana. In the Wye Valley, the history of S. whiteana is somewhat simpler. The recent discoveries stemmed from rope access survey work on the cliffs funded by English Nature. It was found on Ban-y-gor cliffs in 2000 (Houston ef al. 2001), but was not recognised until 2002 after excellent material had been found at Shorn Cliff, Tidenham (Houston et al. 2002). One possible plant was found at Lady Park Wood, Monmouthshire in 2002 but requires con- firmation from fruit (Houston et al. 2003). It has not been found on searches of the other main cliffs on the English side of the Wye Valley (e.g. around Symonds Yat, The Great Doward, Dennis Grove, Wintour’s Leap; SORBUS WHITEANA ql Houston et al. 2003, 2004); large areas of the Welsh side remain to be surveyed. Possibly coincidentally, Salmon (1930) had also cited material of S. hungarica named by T. Hedlund collected from cliffs at Pen Moel and Tidenham in the Wye Valley (v.c. 34). Hedlund’s determination is surprising given that S$. hungarica is part of the S. intermedia group rather than the S. aria group. The S. hungarica folder in Hedlund’s herbarium in UPS contained no British material in 2004, but one sheet, which probably belongs to the S. porrigentiformis aggregate, collected by C. E. Salmon (ref. no. H) on 30 July 1920 from Wye Cliffs, Oakwood, Tidenham had Hedlund’s determination as S. hungarica subsequently crossed out by Hedlund showing that he had later changed his opinion (the duplicate material from Salmon’s gatherings in BM is not S. whiteana either). Hedlund was at the time regarded as an authority on Sorbus, and although he knew the Scandinavian taxa well, he was inconsistent in determination of material from elsewhere in Europe and came to be considered as unreliable (Kovanda 1961 puts it more strongly!). We consider his determinations of British material of S. hungarica as erroneous. Sorbus whiteana qualifies as ‘Endangered’ under the IUCN (2001) Threat Criteria. There appear to be no immediate threats to its survival provided care continues to be taken during conservation habitat management work in the Avon Gorge. Seed from Clifton Down has been deposited in the Millennium Seedbank and plants are being grown at the National Botanic Garden of Wales and _ Bristol University Botanic Garden. ACKNOWLEDGMENTS We would like to thank the Keepers of Herbaria for access to material, English Nature for funding the survey work, John Bailey for counting chromosomes, Philip Oswald for the Latin translation, Ashley Robertson, Colin Charles and Angus Tillotson for help with field work, Clive Stace for numerous improvements to the manuscript, and Naomi and Jenny Rich for helping to extract seeds. REFERENCES AAS, G., MAIER, J., BALTISBERGER, M. & METZGER, S. (1994). Morphology, isozyme variation, cytology and reproduction of hybrids between Sorbus aria (L.) Crantz and S. torminalis (L.) Crantz. Botanica Helvetica 104: 195-214. ALEXANDER, M. P. (1969). Differential staining of aborted and non-aborted pollen. Stain Technology 44: 117- 2, FAY, M. F., GERNANDT, D. S., COWAN, R. S., KITCHEN, M. A. R., KITCHEN, C. & RICH, T. C. G. (2002). Parentage of an unknown member of the Sorbus latifolia (Lam.) Pers. group. Watsonia 24: 91-100. HOUSTON, L., TILLOTSON, A. & CHARLES, C. (2001). Wye Valley Sorbus project. 1. Ban-y-gor rocks Autumn 2000. Unpublished contract report to English Nature, Ledbury, March 2001. HOUSTON, L., TILLOTSON, A. & CHARLES, C. (2002). Wye Valley Sorbus project. 2. Shorn Cliff Autumn 2001. Unpublished contract report to English Nature, Ledbury, March 2002. HOUSTON, L., TILLOTSON, A. & CHARLES, C. (2003). Wye Valley Sorbus project. 3. Upper Wye Gorge Autumn 2002. Unpublished contract report to English Nature, Ledbury, March 2003. HOUSTON, L., TILLOTSON, A. & CHARLES, C. (2004). Wye Valley Sorbus project. 4. Wintour’s Leap and Lady Park Wood Autumn 2003. Unpublished contract report to English Nature, Ledbury, March 2004. IUCN (2001). IUCN Red List Categories. Version 3.1. The World Conservation Union, Gland. KENT, D. H. & ALLEN, D. E. (1984). British and Irish Herbaria. 2nd edition. B.S.B.I., London. KOVANDA, M. (1961). Spontaneous hybrids of Sorbus in Czechoslovakia. Acta Universitatis Carolinae, Biologica 1: 43-83. LEMCHE, E. B. (1999). The origins and interactions of British Sorbus species. PhD thesis, Darwin College, Cambridge. LILJEFORS, A. (1955). Cytological studies in Sorbus. Acta Horti Bergiani 17: 47-113. RICH, T. C. G. & HOUSTON, L. (1996). Sorbus survey of The Gully, Avon Gorge September 1996. Unpublished contract report to English Nature. RICH, T. C. G. & HOUSTON, L. (1997). A new species of Sorbus from the Avon Gorge, Bristol. BSBI News 75: 69. RICH, T. C. G. & HOUSTON, L. (2004). The distribution and population sizes of the rare English endemic Sorbus wilmottiana E. F. Warburg, Wilmott’s Whitebeam (Rosaceae). Watsonia 25: 185-191. ROYAL HORTICULTURAL SOCIETY (1966). R. H. S. colour chart. Royal Horticultural Society, London. SALMON, C. E. (1930). Notes on Sorbus. Journal of Botany 68: 172-177. WARBURG, E. F. & KARPATI, Z. E. (1968). Sorbus, in TUTIN, T. G. et al., eds., Flora Europaea 2: 67-71. Cambridge University Press, Cambridge. (Accepted November 2005) ai pati. ine rege s anh lip sek a > ey Viewer et j Dad Mise itty, tact ber ae ikon ae li nt at ie en i ee va Gs ane ‘eka, ne) 4? . ey a a tia 1.8) sipicnyinain lle nel ee Sha thal: pert i my muni ity ee nie rh aes i eee : Te it. wa + ee Ve tout i hd a ee oe Ate hy * Se. bY all oat ie aaa - al ‘Tole as e ny fa - . 4 ' in Watsonia 26: 9-14 (2006) Sorbus pseudomeinichii, a new endemic Sorbus (Rosaceae) microspecies from Arran, Scotland A. ROBERTSON* School of Biological Sciences, University of Bristol, Woodland Road, Bristol BS8 1UG and C. SYDES Scottish Natural Heritage, 2 Anderson Place, Edinburgh EH6 5NP ABSTRACT Sorbus pseudomeinichii Ashley Robertson sp. nov., a new endemic tree from the Isle of Arran, Scotland is described. Its leaf morphology is intermediate between S. pseudofennica and S. aucuparia and molecular studies support the hypothesis that it was formed by hybridisation between S. pseudofennica and §. aucuparia. All three hybridogenous taxa endemic to Arran, S. arranensis, S. pseudofennica and §. pseudomeinichii, can be _ considered endangered. To maximise their chances of long-term survival, all locations and components of this evolutionary complex should be conserved. KEYWORDS: hybridisation, new species, Sorbus arranensis, Sorbus pseudofennica. INTRODUCTION Recent molecular studies (isozyme, nuclear DNA and chloroplast DNA) have confirmed that the Arran endemics, Sorbus arranensis Hedl. and S. pseudofennica E. F. Warb., are of hybrid origin, the former originating as S. aucuparia L. x S. rupicola (Syme) Hedl., and the latter subsequently as S. arranensis x S. aucuparia (Figure 1; Nelson-Jones et al. 2002; Robertson et al. 2004a). The discovery of an unusual Sorbus specimen, in the herbarium of the Royal Botanic Garden Edinburgh, collected by Dr D. McVean in 1949, prompted Lusby (1996) to suggest that further diversity was being produced within the complex in Arran by crosses between S. pseudofennica and S. aucuparia and, during a detailed study, a small number of trees were found that had a leaf morphology that matched this unusual specimen (Robertson et al. 2004b). Molecular marker analyses support the hypothesis that these trees originated as S. aucuparia x S. *e-mail: ashley.robertson @bristol.ac.uk pseudofennica (Robertson et al. 2004b) and they are treated as a new species for consistency with other hybridogenous Sorbus taxa in Britain. Sorbus pseudomeinichii Ashley Robertson sp. nov. HOLOTYPUS: Glen Catacol, main burn, east bank, Arran (v.c. 100), Scotland, 1 June 2004, A. Robertson (NMW, accession number V.2004.017.21). ISOTYPUS: BM, CGE, E, UPS. Arbor ad 4 m ut minimum. Gemmae lance- olatae, hirsutae. Stipulae leniter dentatae, caducae. Ramunculi juvenes petiolique, ut rhachides, modice tomentosi, pilis albis appressis simplicibus vestiti. Petioli 17-38 mm. Folia 75-130 x 45-105 mm, foliolorum later- alium 4—5 paribus folioque terminali majore, ambitu generali ovata, pinnarum paribus mediis longissimis. Foliola lateralia ad medium foli 25-55 x 9-23 mm, oblonga vel oblongolance- olata, rotundata sed ad basin asymmetrica, ad apicem obtusa vel acuta, marginibus propemodo a basi per unum _longitudinis trientem serratis, dentibus prorsum flexis acuminatis, supra viridia glabrescentia, infra pallidiora et praecipue secus costam venasque sparsim pubescentia; pinnae infimae sessiles vel manifeste petiolatae, superiores saepe ad rhachin margini proximali adnatae. Foliolum terminale foliolis lateralibus plerumque simile sed multo grandius, 32-60 x 15-45 mm, ovatum, in parte media cum vel sine uno duobusve lobis profundis vel non profundis fissum (aliquando usque ad costam, in margine saltem distali, sectum), ad basin cuneatum, ad apicem plerumque obtusum. Pedunculi et inflorescentiae rami tomentosi. Inflorescentia 10 A. ROBERTSON AND C. SYDES : Wa S. aucuparia BB (2n=2x=34) X S. arranensis AAB (2n = 3x = 51) S. pseudofennica AABB (2n = 4x = 68) S. rupicola AAAA (2n = 4x = 68) S. pseudomeinichii ABB ? (2n = 3x = 51) FIGURE 1. Relationships among Arran Sorbus taxa. The letters B and A indicate haploid genomes from S. aucuparia and S. rupicola respectively. Ploidy levels are given in brackets. corymbosa. Sepala c. 1 5—2 mm, triangularia, ad basin sparsim hirsuta, apicem versus glabra, in marginibus glandulosodentata. Petala c. 4 mm, orbicularia breviter unguiculata alba cupulata, in facie superiore sparsim pub- escentia. Stamina c. 20, cremea. Styli 3, usque ad basin discreti. Fructus inter S. pseudo- fennicae et S. aucupariae fructus intermedi. Tree to at least 4 m tall. Buds lanceoloid, hairy. Stipules weakly toothed, deciduous. Young twigs, petioles and rachises moderately tomentose with white, appressed, simple hairs. Petioles 17-38 mm. Leaves 75-130 mm x 45— 105 mm, with 4—5 pairs of lateral leaflets and a larger terminal leaflet, ovate in general outline with the middle pairs of pinnae the longest. Lateral leaflets at middle of leaf 25-55 mm x 9-23 mm, oblong or _ oblong-lanceolate, rounded but asymmetrical at the base, obtuse to acute at the apex, with margins serrate from near the base to about one third of the way along the leaf to the apex, with forward- pointing, acuminate teeth, green and glabrescent above, paler and sparsely pubescent below especially along the midrib and veins; lowest pinnae. sessile or clearly stalked, the upper often adnate to the rachis on the proximal margin. Terminal leaflet generally similar to the lateral leaflets but much larger, 32-60 mm x 15-45 mm, ovate, with or without one or two, deep to shallow lobes at the middle (sometimes cut to the midrib at least on the distal margin), cuneate at the base, usually obtuse at the apex. Peduncles and inflorescence branches tomentose. Inflorescence corymbose. Sepals c. 1 5—2 mm, triangular, sparsely hairy at base and glabrous towards tip, glandular- toothed on margins. Petals c. 4 mm, orbicular with a short claw, white, cupped, sparsely pubescent on upper side. Stamens c. 20, cream. Styles 3, free to base. Fruits intermediate between those of S$. pseudofennica and S. aucuparia. Endemic to Arran, Scotland. Found beside the main burn in Glen Catacol (two plants), and in the tributary Allt nan Calman (one plant). SORBUS PSEUDOMEINICHII 11 tI FIGURE 2. Leaves of Sorbus pseudomeinichii. Scale bar = 1 cm. The altitudinal range is c. 150-200 m. One of the Glen Catacol plants, a sapling when first recorded, was not relocated in 2004 and appears to have died. The population of two trees qualifies it as ‘Critically Endangered’ under the IUCN (2001) Threat Criteria. Sorbus pseudomeinichii (Figure 2) is an example of continuing evolution in Sorbus. It differs from S. aucuparia in having the terminal leaflet larger than the lateral leaflets (the terminal leaflet is about the same size as the lateral leaflets in S. aucuparia). It differs from S. pseudofennica in having largely pinnate leaves with 4-5 pairs of pinnae (typically 1-2 pairs of free pinnae in S. pseudofennica). Sorbus arranensis rarely has any free pinnae and is readily distinguished. The relationships between the taxa are summarised in Figure 1. DISCUSSION Isozyme, nuclear DNA and chloroplast DNA studies have shown that, like S. arranensis and S. pseudofennica, S. pseudomeinichii is a Sorbus hybrid that has ultimately been derived from the genomes of both S. aucuparia and S. rupicola (Robertson et al. 2004b). These studies also show that two of the trees, one of which is thought to have died, shared the same genetic marker profile, implying that they were members of the same apomictic clone. They are genetically distinct from the third tree. Neither of these two genetic marker profiles has been roundebelore im S. arranensis or ‘S. pseudofennica. The simplest explanation for the origin of these S. pseudomeinichii trees is that they are the product of sexual outcrossing between S. pseudofennica and S. aucuparia. This would give a triploid hybrid with a predominance of chromosomes from_ S. aucuparia. The genetic marker evidence is compatible with this hypothesis, as is an examination of pollen grain viability (Table 1), which suggests, in the context of other Sorbus species (Liljefors 1955), that S$. pseudo- meinichii is probably triploid (T. Rich, pers. comm. 2004). The epithet pseudomeinichii has been chosen to draw attention to the similarity in origin to the Scandinavian S$. meinichii (Lindeb.) Hedl., thought to be derived from S. aucuparia x S. hybrida L. (S. fennica Kalm ex Fries) (Liljefors 1953; Salvesen 1992; Bolstad & Salvesen 1999). Liljefors’s (1953) experiments revealed that what was then considered S$. meinichii existed in three forms: a tetraploid apomict (to which the name was originally applied), a triploid apomict and a triploid sexual hybrid. Liljefors (1953) proposed that the name S. meinichii should be retained for the tetraploid and a new name Sorbus teodori Liljef. given to the triploid forms. Bolstad and Salvesen (1999), using morphometric analysis and cytology, studied the relationships between the S. meinichii group, S. hybrida and S. aucuparia. They agreed with Liljefors (1953) that it was likely the S. meinichii forms had been derived from crosses between 5S. aucuparia and §S. hybrida, but disputed the separation of S. teodori from S. meinichii. They showed that S. meinichii and S. teodori are, in reality, at the same ploidy level, which is triploid, not tetraploid as originally thought. 12 A. ROBERTSON AND C. SYDES TABLE 1. POTENTIAL POLLEN VIABILITY ASSESSED USING ALEXANDER’S STAIN FOR ARRAN SORBUS TAXA HELD IN WELSH NATIONAL HERBARIUM (NMW),. DATA COURTESY T. RICH, DECEMBER 2004 Taxon Locality Site code NMW accession % viable No. of grains number pollen counted S. arranensis Diomhan Burn, east bank A V.2004.17.17 2% 85 Diomhan tributary, north bank B V.2004.17.20 0% 100 Catacol Burn, west bank E VEZ 004 Ale 4% i538 Allt Easan Biorach, east bank I V.2004.17.13 5% WS Allt Easan Biorach, east bank M V.2004.17.14 1% 133 Mean 3% S. pseudofennica Diomhan Burn, west bank A V.2004.17.8 64% 60 Diomhan Burn, west bank A V.2004.17.2 716% 105 Diomhan Burn, west bank A V.2004.17.1 68% 89 Diomhan Burn, east bank A V.2004.17.7 83% 96 Catacol Burn, east bank E V.2004.17.3 64% 74 Mean 71% S. pseudomeinichii Catacol Burn, east bank (holotype) C V.2004. 17.21 0% 209 TABLE 2. LOCATIONS AND POPULATION SIZES OF ENDEMIC SORBUS TREES AT 14 SITES ON ARRAN Locality Grid Reference Site Code S. arranensis — §. pseudofennica Most recent Population size Population size Survey Diomhan Burn NR925467 A 209 Dil Robertson 1997 Diomhan tributary NR930467 B 32 23 Robertson 1997 Allt nan Calman NR918454 G 24 33 Robertson 1997 Abhainn Bheag NR924489 D 0 40 Robertson 1997 Catacol Burn NR917457 E 33 . 40 Robertson 1997 Catacol tributary NR917457 E 8 25 Burlison 1986 Creag na h-lolaire NR924475 F 1 0 Burlison 1986 Allt nan Champ NR936430 G 30 0 Burlison 1986 Garbh Coire Dubh NR951432 H 24 0 Burlison 1986 Allt Easan Biorach NR953474 I il 0 Robertson 1997 Allt Easan Biorach NR952467 J 3 0 Robertson 1997 Allt Dubh NR944470 IL, Dah 0 Robertson 1997 Allt Easan Biorach NR945496 M + 0 Robertson 1997 North Glen Sannox NR964457 N 1 0 King 1981 Total 407 436 Previous surveys have identified 14 sites for S. arranensis and S. pseudofennica (Fig. 3), with total population sizes in 1997 of 407 and 436 plants respectively (Table 2). They are restricted to steep-sided, granite gorges on strongly acid soils (Table 3). Site K is no longer populated with either taxon due to a local extinction event, possibly a fire or landslide. Trees were last recorded at this site in 1956. Landslides or fire may also account - for the lack of S. rupicola trees in the area. Competition from other Sorbus taxa or simply that S$. rupicola has never been in the locality are possible alternative explanations. The nearest known stations for S. rupicola are: Gleann Dubh and Holy Island, both Arran and on the mainland, north of Arran, at Ardlamont Point. High mortality rates and a lack of recruit- ment for S. arranensis and S. pseudofennica were used as justifications for the establishment of the Gleann Diomhan National Nature Reserve (McVean 1956; Bignal 1980; Boyd et -al. 1988). A great deal of research has since SORBUS PSEUDOMEINICHII 13 Kilobrannan Sound Lochrartza Glen Catacol FIGURE 3. Locations of Sorbus species on Arran. been carried out on their ecology and conservation, and although successive surveys have indicated an increase in the populations of both species there is no clear evidence to Suggest that successful regeneration is occurring as differences between the survey methods make comparisons difficult (Robertson 2004). A true picture of the population trends will have to await further monitoring, perhaps using a standard protocol as developed for the Sorbus taxa at Craig y Cilau, Wales by Rich (2003). Gleann Easan Biorach 1 km Glen lorsa Understanding the population dynamics of the Arran Sorbus taxa is central to any conservation plan, especially in the short-term. However, it is equally important to plan for the long-term. Long-term conservation plans should be centred on the processes and potential of the taxa to evolve. If evolution 1s thought not to be occurring then conservation policy should be aimed at preserving the unique products of hybridisation that are not to be repeated. If on the other hand evolution can be shown to be ongoing then conservation 14 A. ROBERTSON AND C. SYDES TABLE 3. SOIL TYPES FOR ROOTING ZONES OF SORBUS SPECIES ON ARRAN. SAMPLES WERE AIR DRIED, AND LATER MIXED 50:50 WITH DISTILLED WATER AND PH MEASURED WITH A PHEP2 HANNA POCKET-SIZED PH METER AFTER EQUILIBRATION Taxon Locality S. arranensis Diomhan Burn, east bank Diomhan tributary, north bank Allt Easan Biorach, east bank Allt Easan Biorach, east bank Allt Easan Biorach, east bank Diomhan Burn, west bank Diomhan Burn, west bank Diomhan tributary, south bank Allt nan Calman, north bank Catacol Burn, east bank Catacol Burn, east bank S. pseudofennica S. pseudomeinichii Site Code Soil Mixed black organic and grit, pH 3 5 Mixed black organic and grit, pH 3 5 Black organic peat, pH 3 7 Black organic peat, pH 3 8 Dark grey mixed grit and organic, pH 3 5 Black organic peat, pH 4 1 Dark, organic, pH 3 7 Dark grey mixed grit and organic, pH 3 6 Black organic peat, pH 3 7 Black peat, pH 3 4 Black organic peat, pH 4 0 Mmmawrr fury policy should aim to ensure that management allows this evolutionary process to continue. The data presented here and in Robertson et al. (2004a, 2004b) have shown that the Sorbus complex on Arran is an actively evolving entity with new genotypes being generated through- out the range of the endemic taxa; there are now three Sorbus species that have evolved there. All three endemics can be considered endangered, and to maximise their evolutionary conservation management objective should be to ensure that all the components of the evo- lutionary complex, S. arranensis, S. aucuparia, S. pseudofennica and S. pseudomeinichii are retained in the northern glens of Arran. ACKNOWLEDGMENTS We would like to thank Philip Oswald for the Latin translation and Tim Rich and Michael Proctor for helpful comments on _ the potential and long-term survival, the manuscript. REFERENCES BIGNAL, E. (1980). The endemic whitebeams of North Arran. Glasgow Naturalist 20: 59-64. BOLSTAD, A. M. & SALVESEN, P. H. (1999). Biosystematic studies of Sorbus meinichii (Rosaceae) at Moster, S. Norway. Nordic Journal of Botany 19: 547-559. BoyD, W. E., LAING, A. E., STEVEN, G. & DICKSON, J. H. (1988). The history and present management of the two rare endemic trees on the Island of Arran. Environmental Conservation 15: 65-66. LILJEFORS, A. (1953). Studies on propagation, embryology and pollination in Sorbus. Acta Horti Bergini 16: 277-239. LILJEFORS, A. (1955). Cytological studies in Sorbus. Acta Horti Bergini 17: 47-113. LUSBY, P., WRIGHT, J. & CLARKE, S. J. (1996). Scottish Wild Plants. Edinburgh: The Stationery Office. IUCN (2001). IUCN Red List Categories. Version 3.1. The World Conservation Union, Gland. MCVEAN, D. (1954). Notes on the present position of the endemic Arran Sorbus. Gleann Diomhan Management plan I. NCC144/MP. NELSON-JONES, E. B., BRIGGS, D. & SMITH, A. G. (2002). The origin of intermediate species of the genus Sorbus. Theoretical and Applied Genetics 105: 953-963. RICH, T. C. G. (2003). Establishment of a monitoring regime for Sorbus species at Craig y Cilau NNR. Unpublished contract report to Countryside Council for Wales, January 2003. ROBERTSON, A. (2004). Status review of the Arran endemic whitebeams, Sorbus arranensis and Sorbus pseudofennica. Unpublished report to Scottish Natural Heritage, February 2004. ROBERTSON, A., NEWTON, A. C. & ENNOS, R. A. (2004a). Multiple hybrid origins, genetic diversity and population genetic structure of two endemic Sorbus taxa on the Isle of Arran, Scotland. Molecular Ecology 13: 123-143. ROBERTSON, A., NEWTON, A. C. & ENNOS, R. A. (2004b). Breeding systems and continuing evolution in the endemic Sorbus taxa on Arran. Heredity 93: 487-495. SALVESEN, P. (1992). Sorbus meinichii. Plantsman 13: 193-198. (Accepted November 2005) ! Watsonia 26: 15—25 (2006) Distinction between the sedges Carex vulpina L. and C. otrubae Podp. and the potential for identification of hybrids C. SMITH*! AND P. A. ASHTON*” NGAS Edge Hill College of Higher Education, St. Helens Road, Ormskirk, Lancs. L39 4QP ABSTRACT Carex vulpina and C. otrubae are difficult species to distinguish using floral characters. As an alternative, SEM of the nutlet surface, stomatal morphology and isozymes have been used as taxonomic tools to distinguish between the two taxa. The three methods show differentiation along species lines and are suitable to identify hybridisation. Stomatal morph- ology and isozymes reveal evidence of possible introgression between the two species. KEYWORDS : False Fox-sedge, True Fox-sedge, hybridisation, isozymes, nutlet morphology, SEM, stomata. INTRODUCTION Carex. vulpina L. (Fox sedge) and C. otrubae Podp. (False Fox sedge) have been recognised as distinct species since separation by Ernest Nelmes (1939). Nelmes identified a broad geographical separation between the two species, considering C. vulpina to be a plant predominantly of eastern Europe while C. otrubae is more typical of western Europe. However, both species occur in Britain, with C. vulpina being a rare plant of wet meadows in southern Britain found in only eleven hectads (Preston et al. 2002) while C. otrubae, with a wider ecological tolerance, is more common, being found in a variety of damp habitats. The two species are sympatric at some sites. Hence distinction between the two plants is important to allow the ecology of C. vulpina to be understood and to allow informed conservation. The two species have long been considered difficult to separate due to their morphological similarity and the character variability within the two species. The initial separation (Nelmes 1939) was based upon differences in cell morphology, culm shape, leaf colour and character of the inflorescence (bracts, glumes and utricle shape). Subsequent authors have added to the list of supposedly distinguishing morphological features, Erskine & Lambrick *le-mail: smithco@edgehill.ac.uk *e-mail: ashtonp @edgehill.ac.uk (2000) listed 29 such characteristics. These have typically been qualitative and have included such features as ligule shape (Nelmes 1946), bract shape (Senay 1950) and the timing of utricle fall (Erskine & Lambrick 2000). However Foley & Porter (1999) consider many of these to be of little practical use. Porley (1999), following the pioneering work of Crawford (1910) and Metcalfe (1971) on the distinctiveness of the leaf and stem anatomy of many British Carex species, suggested that internal leaf anatomy could be used to discriminate between the two species, although our preliminary investigations using _ this method have proved inconclusive. Blurring of the delineation between the two species will result if the species hybridise. Nelmes (1939) suggested that hybrids between C. vulpina and C. otrubae had been found close to the River Medway, at Tonbridge (v.c. 16) and from Amberley Wild Brooks (v.c. 13), although Stace (1975) noted that these records are unconfirmed. However, Stace (1975) also considers that “hybrids between the two species will be revealed by careful searching’. In Britain crosses have also been reported between C. vulpina or C. otrubae with members of the Carex muricata group and C. paniculata (Marshall 1897), although further studies of these specimens is required to substantiate their status (Stace 1975). Discrimination between other morpho- logically cryptic Cyperaceae taxa has been undertaken using alternative characters to gross floral morphology. SEM microscopy of the nutlet surface has been successfully used by Schuyler (1971) to distinguish between morphologically similar species of Scirpus and Eriophorum while Flatberg (1972), among other approaches, utilised differences in stomatal measurements to investigate the occurrence of a new hybrid between Carex canescens and C. chordorrhiza in central Norway. Starr & Ford (2001) found that SEM of nutlet micromorphology combined with leaf anatomy and morphology (including stomatal 16 C. SMITH AND P. A. ASHTON TABLE 1A. CAREX VULPINA SITES AND SAMPLE DETAILS Site Map Reference —§ Vice County Isozyme Sample Nos. Stomatal Sample Nos. Amberley Wild Brooks (AMB) TQ038136 vic. 13 1 3 Love Farm East (LF) TQ066276 WAC aus 1 1 Besley Farm Wet Meadow TQ016149 WeSo 3) 1 0 Besley Farm (BF) TQ018149 VAC IS, l D, Blackthorne (BMP) SP636204 WSs 22) 1 1 Otmoor (Nutlets) (OT) SP578128 Wo BS 1 4 Asham Meads (AM) SP590146 Wica 23 0 2 Ashleworth Ham (AH) S$0828261 WHCs 8) 3 5) Hawbridge (HA) SO849277 v.c. 34 1 1 R. Porley (RP) Probably Otmoor Wey 28) 0 i Stuttgart Bot. Garden (GE) 1 5 Czech Republic (CZR) 0 1 Total ie 26 TABLE 1B. CAREX OTRUBAE SITES AND SAMPLE DETAILS Site Map Reference = Vice County Isozyme Sample Nos. Stomatal Sample Nos. Ashleworth Ham (AH) S$O0828216 v.c. 34 1 5 Coombe Hill (CH) SO0873270 v.c. 34 1 2 Point of Ayr (PA) S$J122848 ViCn 0) 1 1 Donington (D) TF210355 VACOS 1 2 Rixton (RI) $J685903 v.c. 59 1 3 Crossens (CR) S$D363214 WAC DY) 1 1 Hesketh Outmarsh S$D425255 V.c. 39 1 0 Gait Barrows (GB) $D418772 v.c. 60 1 1 Port William (PW) NX315488 v.c. 74 1 1 Total 8 16 characters) distinguished between members of Carex section Phyllostachys along traditional taxonomic lines. A number of authors have also used isozymes to address distinctiveness between morphologically similar species (Ford et al. 1991; McClintock & Waterway 1993; Ford et al. 1998; Reinihammar 1999; Tyler 2003). This paper aims to investigate the usefulness of using three of the methods outlined above (SEM of the nutlet surface, stomatal morph- ology and isozymes) to distinguish between C. vulpina and C. otrubae. In addition we comment on the potential of each method for assessing the presence of hybrids. METHODS COLLECTION AND PROPAGATION Species were identified a priori using a range of characters. C. vulpina was distinguished by a stout, sharply winged flowering culm, presence of dark auricles below the inflorescence, the presence of spare, chaffy material around the ligule and the possession of wrinkled leaf sheaths. These are recognised as the most useful diagnostic morphological characters (Rich & Jermy 1998). C. vulpina was collected from eight out of the eleven hectads where it is present in England, (Preston et al. 2002) in addition to two overseas samples ( Table la ). C. otrubae was collected from nine sites across a large part of the range of the species in Britain (Table 1b). At two sites (Amberley Wild Brooks and Ashleworth Ham) the two species were observed growing sympatrically. All of the C. otrubae and most of the C. vulpina material were grown from ramets collected from the various sites. From five C. vulpina locations (Otmoor, Amberley Wild Brooks, Asham Meads, Germany and _ the Czech Republic) nutlets, rather than vegetative material, were collected and subsequently germinated. DISTINCTION BETWEEN CAREX VULPINA AND C. OTRUBAE 17 Successful germination of nutlets was achieved by selecting undamaged, full utricles that were hard to the touch. These were then cleaned of dry debris and stored in paper envelopes at 4 °C. in a jar containing silica gel to ensure a low moisture content. These were collected in late summer/ early autumn. In the following February the utricles were spread on wet filter paper in Petri dishes and kept in the dark at 4 °C. After one month the closed Petri dishes were moved into light in an unheated glasshouse until germination occurred. Throughout these stages drying out was prevented. Within two weeks signs of germination became evident. When the root growth was established and the cotyledons reached about 5 mm in length then the plants were transferred to potting compost in small pots and then grown on outside. Within six weeks from placing the plants in the glasshouse the two species exhibited germination rates between 36-96%. Individuals grown from nutlets were kept until flowering and seed set when identification could be confirmed. Tables showing percentage germination rates for the species in the years 2002 and 2003 are shown in Tables 4a and 4b (see Appendix). SEM METHODS Nutlets were prepared for SEM using a method modified from Starr & Ford (2001). The C. vulpina material was from Ashleworth Ham (v.c. 34) and the C. otrubae material came from Rixton (v.c. 59). The utricles were removed and the nutlets were acetolysed in 1:9 sulphuric acid — acetic anhydride solution using a modified form of Starr & Ford’s protocol. Both sets of nutlets were immersed in the solution and placed in an ultrasonic bath at a temperature of 55 °C. C. otrubae needed an immersion time of 15 minutes whilst C. vulpina required 35 minutes. They were then rinsed in three changes of distilled water and allowed to dry overnight. Two utricles of each species were prepared for scanning. Financial constraints prevented a more extensive sample being taken. STOMATAL MORPHOLOGY Using fresh material from the central section of a mature leaf, microscope slides of he abaxial epidermis were prepared using a method modified from Starr & Ford (2001). This involved immersing the leaf in bleach and scraping away unwanted tissue with a scalpel, rinsing in water, dehydrating in ethanol and staining in 1% safranin prior to mounting. Ten randomly selected stomata were then measured from the prepared slides at 400 x magn- ification. Two measurements were recorded for each stoma, the maximum length, (the overall length of the stomatal complex running co- axially with the stomatal opening) and maximum width, measured across the outer margins of the guard cells (Fig. 1). Width FIGURE 1. Showing the axes from which stomatal measurements were taken. Dividing the maximum length by the maximum width gave the stomatal index (SI). In all cases the stomatal opening was closed. Stomata from 26 individuals of C. vulpina originating from eight English sites plus the two European samples were examined (Tables la and 1b). In the case of C. otrubae stomata from sixteen individuals from eight British sites were measured. Following checking for homogeneity of variances these data were analysed using one-way ANOVA. Post hoc testing was undertaken using the Tukey test. ISOZYME METHODS The two species gave regular consistent banding when assayed on three isozyme systems (glucose-6-phosphate isomerase (GPI) E.C. 5.3.1.9, 6-phosphogluconate dehydro- genase (6-PGD) E.C. 1.1.1.44, and menadione reductase (MNR) E.C. 1.6.99.2.). These isozymes had previously been shown to be useful in addressing taxonomic questions in Carex (e.g. Schell & Waterway 1992; Jonsson & Prentice 2000) and had also been found to exhibit interspecific variation in other Carex species during previous work in our laboratory (Blackstock, Smith & Ashton unpub.). Isozyme methods were modified from Abbott (1993), Ashton (1990) and Soltis (1990). 12% starch gels were used with GPI being run on a discontinuous histidine citrate C. SMITH AND P. A. ASHTON FIGURE 2b. Carex otrubae showing pointed central body and less obvious satellites. DISTINCTION BETWEEN CAREX VULPINA AND C. OTRUBAE + = FIGURE 2d. Carex otrubae elongate surface cells. 20 C. SMITH AND P. A. ASHTON buffer system and 6-PGD and MNR run on a discontinuous tris citrate buffer system (Wendel 1990). Each gel contained samples from both species. The origin of each individual used in each lane is listed in Table 2. RESULTS The two plates, Figs. 2a and 2b, show the differences in the nutlet surface cell morphology between the two species. The terms used to describe the features follow Schuyler (1971). Note the more rounded central body and the presence of smaller, raised satellite bodies offset towards the cell wall in C. vulpina (Fig. 2a). C. otrubae has a more sharply pointed central body and_ less prominent peripheral satellites (Fig. 2b). When viewed perpendicularly surface cells of C. otrubae (Fig. 2d) nutlets are longer along the axis that passes from the stipe to the beak of the nutlet, while in C. vulpina these cells are rather orbicular in outline (Fig. 2c). In all cases the gels were loaded such that lanes 1-11 were C. vulpina ‘and lanes 12-20 were C. otrubae. The three isozyme systems reveal distinction between the two taxa. GPI shows interspecific variation, C. vulpina being monomorphic for the GPI-Ja allele and C. otrubae being monomorphic for the GPI-1b allele (Fig 3a). A similar pattern is observed in 6-PGD, with C. otrubae being monomorphic for the 6-PGD- Ic allele while C. vulpina typically possesses the 6-PGD-Ib allele at this locus (Fig 3b). However two C. vulpina individuals possess a different allele. The specimen from Amberley Wild Brooks (lane 2) having the same allele as that found in C. otrubae (6-PGD-Ic) while the specimen from Hawbridge (lane 8) expressed a unique allele (6-PGD-1a). MNKR reveals a more complex pattern than in the previous isozymes considered (Fig 3c). This has been interpreted as consisting of two loci. C. otrubae expresses only one of these loci (MNR-1) and is monomorphic. The full range of variation is observed in C. vulpina. In this species some individuals have the same pattern as that found in C. otrubae (Amberley Wild Brooks and Hawbridge; tracks 2 and 8 respectively) or they express the slower form of this gene (MNR-Ib, tracks 5 and 6, Ashleworth Ham). Alternatively the MNR-1 band is absent and individuals exhibit either a single MNR-2a band (lanes 9 and 11; Stuttgart and Besley Wet Meadow) or a two- banded pattern combining either MNR-2b and 2c (lanes 1, 4 and 7; Blackthorne, Ashleworth Ham and Love Farm) or MNR-2a and 2b (Lanes 3 and 10; Ashleworth Ham and Besley Farm). There is considerable overlap in mean stomatal length in C. vulpina and C. otrubae and some overlap in stomatal width (Table 3). Stomatal shape differences between the two species are clearest when both measurements are considered together (Fig. 4). C. otrubae stomata are wider than they are long (below the diagonal line in Fig. 4) having a stomatal index (SI) of less than 1. By comparison C. vulpina has stomata that are above the diagonal line in Fig. 4, being longer than they are wide (SI > 1). When the mean values of all populations, when n = 10, are considered using one-way Anova there is a_ statistically significant division (F 47432= 3997, P< 005). The only sample which does not follow this pattern is one of the C. otrubae specimens from Ashleworth Ham, which clusters with C. vulpina. SI values for each individual surveyed are given in Tables 5a and 5b (in appendix). DISCUSSION Discrimination between the two species can be achieved using stomatal shape, isozyme pattern and nutlet micromorphology. Within the small sample used for SEM there is consistency within a species and a difference between the two species. C. otrubae has a more sharply pointed central body with few, if any, peripheral satellite bodies. C. vulpina displays the outer satellite bodies with a blunter central body. Nutlet surface cells in C. vulpina tend to be orbicular in outline while C. otrubae cells have a pronounced longer axis in a line extending from the distal to the proximal end of the nutlet. Similarly when stomatal shape is compared C. vulpina has elliptical (sub- orbicular) stomata while C. otrubae tends to have rounder stomata (orbicular). The differences in isozyme pattern are slightly less clear cut. The interspecific variation coupled with the absence of intraspecific variation renders GPI the most useful system for delineating between the two taxa. However the patterns observed in 6-PGD and MNR are almost as informative. Despite the apparent complexity of MNR in C. vulpina, only 20% of C. vulpina expressed the same pattern as that found in C. otrubae. 21 DISTINCTION BETWEEN CAREX VULPINA AND C. OTRUBAE UNI FINAZNA AHL YOd WVADOWAZ 9 TANI aueq0; | Gk) Sk) LES St ST OPEL USE CL OT, OF 8 L 9 S r € Z I 97-UNW = [eal ra q7-UNW mo = — pa 07-YNW — = . res qI-UNW — = DI-UNW @ al Ga (ea) Pay fe] pa = =a] | al ‘GDd-9 ANAZNA AHL YOU WVYADOWAZ Ae FAN auEy 0% 61 St Lk oF SE tl eh °leaf width/ <= leaf width Female glume colour Female spike (distal) orientation - erect/pendent Female spike (distal) male at apex - absent/present Female spike (proximal) orientation - erect/pendent Utricle beak - indistinct/distinct short-beak Utricle nerves - absent/present the leaves of grasses (Poaceae) (Rice ef al. 1979). Stomata were counted from near the central vascular bundle of the leaf in a quadrat formed by a square graticule using 400x magnification. Five replicates per surface were counted per specimen, and counts were con- verted to density/mm”. It is intended that voucher specimens will be deposited at Liverpool Museum (LIV) at the end of the associated work. RESULTS MORPHOLOGY Eleven of the 17 qualitative traits considered are shared by C. recta and C. acuta, four do not present consistent traits and only two can be used to differentiate between the two species (Table 3). This lack of variation precluded any meaningful form of multivariate analysis of the qualitative data. The two are angle texture at the top of the culm (smooth in C. recta and rough in C. acuta), and absence/presence of nerves on the utricles (absent in C. recta and present in C. acuta). Comparing the traits presented by the Bonar Bridge sample for these M. DEAN AND P. A. ASHTON 34 Z IQR, Ul UMOYS se oIe S}TeI) JoJORIeYO puL sidjORIeYD PlOg Ur UMOYs oe soroads oy) UDIMJOq YSINSUNSIp Jey) Sso}OBILYO OM) OU, DJIA“) OOT yuosqe qoq OO! poxeoq-oys yoq C6 100.10 e/U LG juasqe {oq OOI ooo qo0q 00 yoeylq / ep DIOL ) CL YPM feo] < DIAL “J OOI oynoe oq O01 SsoT[Aqr 10u yjoq OOI IAROUOD yoq OOT snouosiy you yjoq Z9 ayeord Apyeam DINID “yD se jou ZR juasqe 4joq c8 juasqe ByNIV “) WLlE “PIII %6H9 ~=69 YyOouUs qjoq OOI oyNIe y0q 001 juosoid JUSLUUSI[e Op ye.) UOXe) dSplLig JeuOg ou OO yuoso.id 16 quosqe SALOU BPLN | Soh OO! poyxeoq-J10Ys OO! poxeoq-joys yeoq sfolyn =O] sok OO! 010 L8 yoo010 uonejustIo xo1d ‘ds } Cy dAISN[OUOOUT Cg juosoid €S yuasqe xode gyeur ysip ‘ds yp] sok OO! 010 L6 ee) e) uo UoLIO \sIp “ds JE] sok 001 poriq/ ep = OOT = RIG. / Jaep IMOJOS ouIN]s JZ] [enaed OL Ypim fesp=Io> [9 YIPIM Jeol < sug] oAHepes gnsiy | | jenaed OS oynoe L6 oynoe odeys xode aginst, (| So OO! dSO][Lql JOU OO! ISO[[UGL Jou SuUIAKIOp UO'YIYS Jeoy 6 Sok C9 JAROUOD 146 JARNUOD odeys ynoury Ys Jeoy sok OOT snouost} JOU C6 snouosLy JOU adeys dy yeay oA sok (8 ayeord Apyeam 99 = ayeord Apyeom ad} UONOOS Jeo] BDA jensed OOT juosoid (S(e juosqe Yj99} UISIeU Jeol BOA CC sok C6 juosqe PL yuasqe yoo) UISIeU Jeo] Wynd =p ou OO ysno.d 06 YyJoous 9.1N}X9)} ojsue WHND = ¢ SoA OOT aynoe OOT oynoe adeys ojsue wynd =Z SoA OO] juosoid 6 yuasoid sojoume =| ye.) poreys OA Wey Wey Joyorieyy) “ON pInIvD “yd DIIAA “Dd DINID “Dy ®W vjJIas “DY SLIVUL YALOVUVHDO AALLVLITVNO NOXV.L FOCI UVNO” ANV VIAOV XAYVO ‘VIOTY XTAVO '¢ ATAVL CAREX RECTA 35 TABLE 4. QUANTITATIVE CHARACTER MEANS (OUTLIERS INCLUDED) FOR CAREX RECTA, CAREX ACUTA AND BONAR BRIDGE TAXON Character C. recta C. acuta Bonar Bridge je Proximal bract length/mm 254 a 216b Zi ste Proximal bract width at midpoint/mm 62a 41b AC caer Culm leaf length/mm 538 a 738 b 507 a ee Culm leaf width at midpoint/mm 67a 5.3) 19 5)) 1) ots Culm length/mm 728 a 1075 b 640 ¢c ta Inflorescence length/mm 181 ab 191 b 167 a a Leaf (vegetative) length/mm 1047 a 1243 b 1001 a Eee Leaf (vegetative) width at midpoint/mm 59a 66b 54a te Ligule length/mm 65 D7 65 ns Female spike length/mm (distal female spike) 322a 48 2b ala ts Peduncle length/mm (distal female spike) 45a 04b 5) a wt Female spike length/mm (proximal female spike) 60 58 5 ns Peduncle length/mm (proximal female spike) 142a 3 6b 20 0c pee Male spike length/mm (terminal spike) 300a 455b 26 55C eh P = level of significant difference: *<0 05; ** <0 01; *** 90%. The second DA of the two even sized groups of 20 specimens and using eight characters also correctly classified each specimen as C. recta or C. acuta. The predicted classification for the Bonar Bridge specimens in the second DA was 38 C. recta and one C. acuta. 34 of the 38 are classified with 100% probability, and the remaining four with probability >95%. The one specimen classified as C. acuta was one classified as C. recta with <100% probability in the original analysis. STOMATAL DISTRIBUTION AND DENSITY There are differences in stomatal distribution between the C. recta samples. The Wick sample has no or few stomata on its adaxial surface whereas the other two samples are amphistomatous (Table 6). C. acuta has abaxial stomata and the abaxial stomatal density is much higher in this taxon than the other two taxa. The Bonar Bridge sample is amphi- stomatous and presents minimum, median and maximum densities that are similar to those of the C. recta samples from Beauly and Invershin. DISCUSSION Before the taxonomic position of the Bonar Budge — taxon ‘can be addressed, the morphological characters that differentiate between C. recta and C. acuta need to be confirmed. Qualitative data showed that only two of the characters studied, smooth or rough culm angle texture at top of culm, and nerveless or nerved utricles, separated C. recta and C. acuta respectively (Table 3). To use just these two characters is insufficient, and is not completely reliable as some C. recta speci- mens had rough culm angles, so quantitative characters also need to be included. However, most of these show considerable overlap, even if the means are significantly different (Table 4). This emphasises the difficulties in working with morphologically variable taxa. It may not be possible to identify an individual specimen with certainty if it is not of typical form. Therefore it is beneficial to measure characters from sufficient specimens selected randomly from recognised populations to produce a table of character means, against which new specimens can be assessed. Considering the characters that may be useful in separating the two species, in general C. recta has shorter culms and culm leaves, shorter distal female spike and terminal male spike, and longer peduncles. In contrast, C. acuta is a larger sedge, with longer terminal male spike and distal female spike, but short peduncles, the distal female spike usually sessile. In addition, with sufficient specimens multivariate analyses can be used, and both the DA with the full C. recta sample and the DA with the equal sized C. recta and C. acuta samples classified all C. recta and C. acuta specimens correctly. This confirms that the set of characters used can differentiate between C. recta and C. acuta. A hybrid taxon may be expected to be morphologically intermediate between the parental species in most characters (Flatberg 1972; Waterway 1994). However, an inter- mediate morphology is not found in the Bonar Bridge samples. Comparison with the differ- entiating criteria for qualitative characters, the Bonar Bridge sample usually has smooth culm angles and nerveless utricles, indicating an affinity with C. recta (Table 3). Analysis of quantitative characters (Table 4) indicates the Bonar Bridge samples generally have lower means than the other two taxa. Except for peduncle lengths, the Bonar Bridge sample is consistent with a smaller form of the C. recta found elsewhere in the British Isles. The 38 M. DEAN AND P. A. ASHTON sample means for several of the length characters, culm leaf, culm, inflorescence, vegetative leaf, distal female spike and terminal male spike, are significantly lower than the C. acuta sample, so the Bonar Bridge sample is much smaller than C. acuta. Overall, of the three taxa, the Bonar Bridge sedges are the smallest. Stomatal density and distribution also align the Bonar Bridge samples with C. recta (Table 6). PCA did not separate the three taxa, but the complete overlap of the Bonar Bridge and C. recta samples along the first two axes implies a very close similarity between the two (Fig. 1). DA also identified a close proximity between C. recta and the Bonar Bridge taxon, class- ifying all 39 specimens as C. recta in the first run, and all but one in the second run with reduced volume of data. There is also strong circumstantial evidence against the presence of a C. acuta x recta hybrid at Bonar Bridge. Although pre-1970 records indicate that C. acuta was found at Strathpeffer in the same vice county as the presumed hybrid (Duncan 1980), the current distribution of C. acuta in the UK extends no further north in Scotland than the Clyde-Forth line, so the recognised distributions of C. acuta and C. recta do not overlap (Preston ef al. 2002). In addition studies including exper- imental hybridisation between C. recta and C. acuta produced a low mean seed set, although this was based on a small sample (Faulkner 1973). Combining these two observations it seems unlikely that this inter-specific cross would have occurred naturally. The evidence presented from morphology, stomatal distribution and geographical location Support the reassessment of the Bonar Bridge taxon as C. recta. This population and other small ones down river are new records for C. recta in East Ross-shire (v.c. 106). This increases the distribution of the plant in the UK from three to five 10-km squares. The C. recta population at Invershin, approximately 5 km upriver, is proposed as the progenitor of this population. Clumps detached from the main sedge stand at Invershin have been observed at the edge of the river, and could float downriver and be left stranded on the shoreline or on the edges of the creeks. It is proposed that such events led to the founding of the Bonar Bridge populations and the other small populations found along the estuary. The Bonar Bridge site to the east of the road bridge is very exposed, resulting in a smaller stature, and this may be a product of phenotypic plasticity. CURRENT DISTRIBUTION EASTERNESS V.C. 96 On the north bank of the River Beauly approximately | km east of the town of Beauly at NH539465. There is a large stand on the flat area beside the river and below the river defence bank. There are scattered sedges in the lagoon area just north of this at NH539466, but these become hidden by the tall, dense vegetation of Phragmites australis during the summer. EAST ROSS-SHIRE V.C. 106 There are three locations on the south-western bank of the Kyle of Sutherland. Furthest upriver near Kilmachalmack from NH503990 to NH508989 although this stand is not continuous. The second location is below Carbisdale Castle, where there are stands in marshy areas beside the river at NH573958, and there is a large stand of C. recta on the riverbank just upriver from the railway bridge at NH578953. The third location starts at Bonar Bridge at NH6091. There are small stands to the west of the road bridge and one large population to the east of the road bridge. There are several small populations along the river’s edge downriver as far as Kincardine including stands at the following grid references: NH604910, NH607894, NH607896 and the furthest down- river at NH605898. EAST SUTHERLAND V.C. 107 There are locations on the north-eastern bank of the Kyle of Sutherland opposite those on the south-western bank in v.c. 106. Furthest upriver near Altass there is a stand on the riverbank at NH504992 and two stands in a marsh at NH503992 and NH5059972. At the second location near Invershin the population extends along the riverbank from NH575960 to NH578956. Some specimens show signs of hybridisation, probably with C. nigra (L.) Reichard. Small, scattered stands of C. recta are found upriver at the heads of small bays, for example at NH571967, NH570967 and NH570968. Only one small stand has been found at the third location, Bonar Bridge, at NH610914. CAITHNESS V.C. 109 On both banks of the River Wick, but mainly on the north bank, just upriver from the town of Wick. On the north bank there is a large stand at ND354514. There are stands extending from ND354514 to ND350516, but amongst these ———— CAREX RECTA 39 there are stands with specimens that are introgressed or have hybridised with C. aquatilis Wahlenb. On the south bank the stands of C. recta show morphological char- acters of introgression to C. aquatilis. There is a stand at ND353513, and there are other small stands close by. ACKNOWLEDGMENTS We would like to thank Brian and Barbara Ballinger for bringing to our attention the stands of sedges at Bonar Bridge and for considerable help with fieldwork. The authors would also like to thank the vice county recorders Margaret Barron, Ken Butler, Dave Earl, Jean Green, Geoffrey Halliday, Morven Murray and Peter Wortham, and Scottish Natural Heritage, Highland Ranger Service, North Wales Wildlife Trust and Sefton Council for help in locating sedge populations or providing access. Thanks also to OXF and MANC for access to their facilities. This work has been undertaken at Edge Hill as part of a research studentship. REFERENCES BLACKSTOCK, N. & ASHTON, P. A. (2001). A re-assessment of the putative Carex flava agg. (Cyperaceae) hybrids at Malham Tarn (v.c. 64): A morphometric analysis. Watsonia 23: 505-516. CAYOUETTE, J. & CATLING, P. M. (1992). Hybridization in the Genus Carex with special reference to North America. The Botanical Review 58: 351-440. DUNCAN, U. K. (1980). Flora of East Ross-shire. Botanical Society of Edinburgh, Edinburgh. FAULKNER, J. S. (1973). Experimental hybridization of north-west European species in Carex section Acutae (Cyperaceae). Botanical Journal of the Linnaean Society 67: 233-253. FLATBERG, K. [. (1972). Carex x lidii Flatb. = C. canescens L. x chordorrhiza Ehrh., a new hybrid. Norwegian Journal of Botany 19: 91-106. HAIR, J. F., ANDERSON, R. E., TATHAM, R. L. & BLACK, W. C. (1998). Multivariate Data Analysis. Prentice- Hall International (UK) Ltd, London. HYLANDER, N. (1966). Nordisk Kdrlvdxtflora. Almqvvist & Wiksell, Stockholm. JERMY, A. C., CHATER, A. O. & DAVID, R. W. (1982). Sedges of the British Isles, 2nd ed. Botanical Society of the British Isles, London. MARREN, P. (1999). Britain’s Rare Flowers. T. & A. D. Poyser Ltd, London. MCCLINTOCK, K. A. & WATERWAY, M. J. (1994). Genetic differentiation between Carex lasiocarpa and C. pellita (Cyperaceae) in North America. American Journal of Botany 81: 224-231. MINITAB INC. (2001). MINITAB User’s Guide. USA. PRESTON, C. D., PEARMAN, D. A. & DINES, T. D. (2002). New Atlas of the British and Irish Flora. Oxford University Press, Oxford. QUINN, G. P. & KEOUGH, M. J. (2002). Experimental Design and Data Analysis for Biologists. Cambridge University Press, Cambridge. EPKA, R. (2003). The Carex muricata aggregate in the Czech Republic: multivariate analysis of quantitative morphological characters. Preslia 75: 233-248. RICE, J. S., GLENN, E. M. & QUISENBERRY, V. L. (1979). A rapid method for obtaining leaf impressions in grasses. Agronomy Journal 71: 894-896. ROSE, F. (1989). Grasses, Sedges, Rushes and Ferns of the British Isles and north-western Europe. Penguin Group, London. SELL, P. & MURRELL, G. (1996). Flora of Great Britain. Butomaceae-Orchidaceae. Cambridge University Press, Cambridge. STACE, C. A. (1997). New flora of the British Isles, 2nd ed. Cambridge University Press, Cambridge. STANDLEY, L. A. (1987). Taxonomy of the Carex lenticularis complex in eastern North America. Canadian Journal of Botany 65: 673-686. SYLVEN, N. (1963). Det Skandinaviska flora-omradets Carices distigmaticae. Opera Botanica A Societate Botanica Lundensi, Botaniskea Notiser 8: 1-161. WATERWAY, M. J. (1994). Evidence for the hybrid origin of Carex knieskernii with comments on hybridization in the genus Carex (Cyperaceae). Canadian Journal of Botany. 72: 860-871. WIGGINTON, M. J. (1999). British Red Data Books: I Vascular Plants, 3rd ed. Joint Nature Conservation Committee, Peterborough. (Accepted October 2005) * 7 \ ce £1 oa was 4 Sah ot ee i ira a bai in he Pow Sey 4 ne ue eye : 4 ‘ , ‘ i - ” - bad . - = - Watsonia 26: 41-49 (2006) 4] Changes in the floristic composition of sand-dune slacks over a twenty-year period P. H. SMITH* 9 Hayward Court, Watchyard Lane, Formby, Liverpool L37 3QP ABSTRACT The aim of this study was to compare vascular plant species-richness and composition in 26 dune-slacks in Birkdale Sandhills, Merseyside during 2003 with the results of a similar survey conducted in 1983. A total of 213 taxa was recorded, as opposed to 150 in 1983, a 42% increase. There is a significant positive correlation between the number of taxa per slack in the two surveys and a significant relationship bet- ween species-richness and slack area in both years. Thirty-six notable taxa were recorded in 2003, an 80% increase on the 20 found in 1983. Fifteen taxa have undergone major increases in frequency, while seven, particularly maritime species, show major declines. The reasons for these changes are discussed and recommendations made for the conservation of the slack flora. KEYWORDS: compositional change, conservation, dune-slacks, notable taxa, species richness. INTRODUCTION In 1983, Sefton Metropolitan Borough Council commissioned a report from the Lancashire Trust for Nature Conservation (now the Lancashire Wildlife Trust) on the ecology of the Birkdale frontal dunes, a narrow strip of sand-dunes 4-23 km long and 120 to 215 m wide, west of the coastal road between Ainsdale and Birkdale (SD322164—SD300122). This dune area was subsequently added to the Birkdale Sandhills Local Nature Reserve. The report (Smith 1983) drew particular attention to the slacks within the study area. These were considered to have special conser- vation interest, being rich in vascular plants, including many uncommon or rare species. In addition, several of the slacks were used for breeding by the endangered Natterjack Toad (Bufo calamita). Having mapped the study area using aerial photographs, the 26 slacks present were searched for vascular plants in June and July *e-mail: phil@smith1941.fslife.co.uk 1983, the richest supporting 78 taxa. Fifteen slacks were considered to have an “incipient” character; i.e. recently created by wind-erosion and therefore in an early stage of vegetation development. Indeed, reference to a time sequence of aerial photographs showed that many of the slacks had formed during the exceptionally dry period of the mid-1970s. Twenty years on from the original study, it was decided to resurvey the slacks, using similar methodology, to determine how species rich- ness and composition had changed and to make recommendations for management. In addition to its Local Nature Reserve status, the study area lies within the Sefton Coast SSSI, a candidate Special Area of Conservation and a Ramsar site. METHODS Each of the 26 mapped slacks (Fig. 1) was revisited at least twice in both June and July 2003. Search time was roughly in proportion to the size of the slack but averaged about 30 minutes per slack per visit. The presence of all vascular taxa (species, sub-species and hybrids) was recorded. Care was taken to include only plants associated with the slack floor and not the surrounding areas relatively unaffected by the water-table. An assessment of slack type was made using the water-table criteria of Ranwell (1972), and the need for management was determined. Slack areas were measured using the Sefton Coast Geographic Information System (GIS). RESULTS SLACK HABITAT | There was no evidence for changes in slack areas, such as infill by blown sand, between the two study periods. The 26 slacks have a total area of 3-31 ha, a size range of 93 to 9856 m’, 42 P. H. SMITH £40 Sia 32 P35" 39-933 534 43 C744 c & g & brs a 02 : (; 1H Sands Lake FIGURE |. Map of Birkdale frontal dune slacks. and a mean area of 1274 m’. Only one slack (no. 44; 875 m’) was considered to be incipient, having a relatively young open vegetation type, compared with 15 slacks in 1983. Using Ranwell’s (1972) water-table criteria, 16 slacks were classed as wet-slacks, six as dry-slacks and four as semi-aquatic (Table 1). Twenty- five slacks have a “mature” vegetation type characterised by a near complete plant cover and a high frequency of shrubs, especially Salix repens. In addition, eight of the wet-slacks have a semi-aquatic component due either to scrapes having been dug for Natterjack Toad breeding pools since the first study or to the presence of an open drain outfall (slack no. 27). SPECIES-RICHNESS A total of 213 vascular taxa was recorded in the 26 slacks, compared with 150 in 1983, a 42% increase in 20 years. Only 22 taxa seen in 1983 were not refound in 2003, representing 15% of the 1983 flora. The range of taxa per slack was 32 to 95 in 2003, as opposed to 4 to 78 in 1983 (Table 1). Mean numbers per slack were 60 3 in 2003 and 41 8 in 1983. This difference is statistically highly significant (t = 607; p = 00001). On average, species-richness has increased by 18 7 taxa per slack. There is a Statistically significantly positive relationship between taxon-number per slack in 1983 and 2003 (r = 0595; p = 0 001) (Fig. 2); FLORISTIC COMPOSITION OF SAND-SUNE SLACKS 43 TABLE 1. SEACK AREAS, HABITAT AND SPECIES-RICHNESS Slack no. (N. to S.) Area (m*) Slack habitat 1983 No. of taxa Slack habitat 2003 No. of taxa 30 1924 Dry 44 Dry 50 31 694 Wet oy Mature wet 51 32 393 Wet incipient 18 Dry Sy 38 354 Wet incipient 37] Dry 56 34 124 Wet Sill Mature wet 39) 35 660 Wet incipient 34 Dry 52 36 105. = —Dry 4] Dry 56 29 93 Wet +4 Mature wet 34 37 1649 Dry 53 Mature wet 68 38 325 _ Wet incipient 16 Semi-aquatic 50 28 2001 Wet 51 Mature wet/Semi-aquatic 86 Dy 2063 Wet 78 Mature wet/Semi-aquatic 95 39 9856 Semi-aquatic incipient 64 Semi-aquatic 93 40 462 Wet 28 Mature wet 56 4] 718 Wet 35 Semi-aquatic 59 42 6625 > Dry; 39 Dry 52 43 1021 Wet incipient 30 Mature wet/Semi-aquatic 15 44 875 Wet incipient - Wet incipient a. 45 . 372 Wet incipient A7 Mature wet 84 46 702 Semi-aquatic incipient 46 Semi-aquatic 51 47 1327 Wet incipient 47 Mature wet/Semi-aquatic 74 48 636 Wet incipient 54 Mature wet 59 49 1988 Wet incipient 54 Mature wet 65 50 2523 Wet incipient 54 Mature wet/Semi-aquatic 66 51 1240 Wet incipient 38 Mature wet 56 Sy 365 Wet incipient 48 Mature wet 70 100 — 8 a @ 80 70 60 tye) =e) N 40 | 30 + 20 5 10 - 0 - 0 20 40 60 80 100 1983 FIGURE 2. Number of taxa per slack, 1983 vs. 2003 (r = 0 595; p =0 027). 44 P. H. SMITH 0 500 1000 Number of species & Ol j=) oO BF @ | 1500 2000 2500 3000 Slack area (m2) FIGURE 3. The relationship between number of taxa and slack area in 1983 (slack no. 39 excluded) (r = 0-416; p = 0-020). 100 90 | — 80 TOG 60 50 No. of species 40 | 30 + 20 0 500 1000 2000 2500 1500 3000 Slack area (m2) FIGURE 4. The relationship between number of taxa and slack area in 2003 (slack no. 39 excluded) (r = 0-399; p = 0-027). in other words, the most species-rich slacks in 1983 tended also to be the richest in 2003. Thus, the slack with the highest number of taxa is no. 27 with 78 plants in 1983 and 95 in 2003. Similarly, the incipient slack no. 44 with only four taxa in1983, had 44 by 2003, one of the lowest counts in the second survey (Table 1). However, this relationship may be affected by other factors, such as slack area. It could reasonably be expected that larger slacks would support more taxa. Regression analysis shows that the relationship between species-richness and slack area is positive and significant in both years (1983: r = 0-416; p = 0-020; 2003: r = 0-399; p = 0-027). Clearly though, this . relationship is weak (Fig. 3, Fig. 4). The r eS, OLS a OY FLORISTIC COMPOSITION OF SAND-SUNE SLACKS 45 TABLE 2. NOTABLE VASCULAR TAXA IN THE BIRKDALE SLACKS Occurrences per slack Taxon 1983 Anacamptis pyramidalis Anagallis tenella Apium inundatum Baldellia ranunculoides Blysmus compressus Carex disticha 1 Carex pseudocyperus Carex viridula subsp. viridula 11 Catabrosa aquatica Centaurium littorale 10 Centaurium pulchellum 3) Dactylorhiza incarnata subsp. coccinea 19 Dactylorhiza purpurella 1 Epipactis dunensis 1 Epipactis palustris 14 Epipactis phyllanthes Equisetum variegatum 9 Hippuris vulgaris Juncus ambiguus Juncus balticus 6 Juncus maritimus 3 Myosotis ramosissima Oenanthe fistulosa Oenanthe lachenalii Parnassia palustris + Potamogeton gramineus Potamogeton pusillus Pyrola rotundifolia subsp. maritima + Salix xangusensis Salix xfriesiana fel Samolus valerandi Dy Schoenoplectus pungens Schoenoplectus tabernaemontani 3 Trifolium fragiferum 10 Triglochin palustris 5 Vulpia fasciculata 1 Status 2003 Nationally Nationally Species of Conservation Rare Scarce Importance 3 3 D 1 2 + 2 + 10 + 2 + 1 + 6 + 1 + 13 + + + + 23 + + 3 + 6 + + 22 + 1 +r

70%) of the wool that the Bradford company scours comes from UK or Irish clips, the remainder from Norway and EC countries such as France, Germany and Holland. Thus, there has been a trend towards importing clean wool, a greater reliance on UK and Irish raw wool and a concomitant decline in the importation of raw wool for scouring by a few specialised companies. As a result, the waste from the woollen processing and manufacturing industry of predominantly foreign origin has become a rare commodity in the past twenty years. There has thus been a considerable reduction in the potential wool alien seed bank originating from regions outside Great Britain and Ireland. The nature of the shoddy industry has changed considerably in the past twenty years. Few of the ten textile waste processors in the Kirklees and Wakefield Municipal Boroughs of West Yorkshire import rags from abroad and if they do, European Community regulations are such that the rags must be cleaned before import. Most rely on UK sources for c.90% of their manufacturing base, acquired either via high street charities or from the textile waste reclamation banks of local authorities. The proportion of woollen to synthetic rags in such consignments is usually low and wool raw materials far less common, so that inevitably, a greater proportion of the processed shoddy fibre for spinning is synthetic. Whether wool or synthetic rags are used, there is little waste and this merely in the form of zips, studs, buttons and labels, which finds its way to a skip and thence a landfill site. The UK woollen manu- facturing industry in general is in decline, in the face of economic market forces relative to European Community and Third World trade agreements. Take for example, the three mills at Kirkheaton, Huddersfield, the heirs of the Jarmaine Company, whose wool waste dump was visited by both John Dony and Ted Lousley and from where Kit Rob and David McLintock first recorded Medicago laciniata in 1955 (Wilmore 2000). One has changed its operation to merchandising of cloth, a second was placed in administration in late 2004 and the third changed to cotton production and closed in March 2005. CHANGES IN AGRICULTURAL PRACTICE AND ENVIRONMENTAL FACTORS The distribution of wool aliens, particularly in West Yorkshire, is now largely dependent upon the agricultural practices associated with the production of a particular vegetable crop, espe- cially rhubarb, winter cauliflowers and spring cabbage. The waste is spread on the land in May-June, ploughed into the host soil in autumn and crowns of rhubarb planted, there to remain for two -years, until transfer to the forcing sheds in the third autumn. In these two growing seasons, the casuals germinate and some apparently set viable seed. Once the crowns have been transferred, the land is again treated with wool waste, ploughed and set down to winter cauliflowers followed by spring cabbage or wheat for a further two or three years. The slow release of nitrogen from the wool waste allows for the production of two or three crops from one spread. The amount of wool waste required per acre is only 25% the volume compared with farmyard manure, which has to be applied annually and is much more difficult to spread. The various types of woollen waste vary in terms of their ability to retain moisture as a mulch and in their nitrogen concentration. They all are in the top group of 130 D. W. SHIMWELL organic nitrogen sources, being ranked with guano and hide meal at 10%, with only hoof and horn meal of a higher concentration at 12% (see Table 2). Precise data is lacking on the relative nitrogen contents of the various types of wool waste, but the noils produced by the scouring process seem to have a particularly high concentration. This encourages the growth of a weed flora dominated by nitrophilous species such as Chenopodium album, Galium aparine and Urtica urens, and rather unusually, Impatiens glandulifera, a prominent and persistent weed of rhubarb fields. The gregarious nature of such plants tends to restrict the growth of other less competitive, annual wool casuals, many of which are denizens of drier soils typical of, for example, Mediterranean Europe. In the Rhubarb Triangle the predominant soil type is a clayey or loamy pelo-stagnogley of the Dale Association, a type of comparatively restricted distribution overlying Carboniferous shales in scattered areas from Sheffield in South Yorkshire, north to Ilkley (Soil Survey of England and Wales 1977, 1983). Typical brown earths of the Rivington | association also occur over Carb- oniferous sandstones, but both types are so modified by fertiliser addition as to restrict natural occurrences to a few narrow strips along a network of footpaths, old railway banks and narrow field boundaries. Many casuals of the Cardueae Tribe of the Asteraceae appear to have suffered due to the progressive reduction in marginal habitats on natural soils and in their inability to withstand competition from gregarious weed species. This group includes species of Centaurea such as C. calcitrapa, C. solstitialis, C. melitensis, C. diluta, along with Carthamus tinctorius, C. lanatus, Silybum marianum and Scolymus maculatus. An increase in a highly competitive nitrophilous weed flora thus appears to have further restricted the frequency of opportunity for annual and biennial wool aliens typical of relatively xeric soil conditions. Rhubarb was first grown in the Wakefield region of West Yorkshire in the late-nineteenth century and by 1938, the 3818 acres accounted for 52:8% of the total acreage of the crop in England and Wales (Beaver 1941). The concentration was truly remarkable and a vital mainstay of many small farms; in the parish of West Ardsley 40% of the arable land was devoted to the crop, in Morley parish 36%, and in Outwood 34%. Beaver remarks: “The soils are not by nature rich, and intensive cultivation can only be sustained by frequent heavy applications of manure — which, incidentally, is now less easy to obtain than in the days when horse-drawn drays and not lorries were the chief means of transport in the towns.’ The implication here is that the use of wool waste had not yet become the predominant method of fertilisation in the late-1930s. This is confirmed by local information that describes the acquisition in 1941-42 of a new rhubarb variety ‘Timperley Early’, from the eponymous Cheshire village, where it was grown with great success by the application of liberal amounts of sewage and night soil. The same treatment was tried in the West Riding for a few seasons, but gradually the almost exclusive use of readily available wool waste came to predominate in the late forties. In the same decade, the market gardeners of Bedfordshire began to import large quantities of wool waste, but other vegetable producing regions such as_ the peatlands of Lincolnshire, Cambridge and south Lancashire continued to rely upon sewage, night soil and farmyard manure. Since the maximal extent of the industry in the late 1930s, a variety of market forces and agri- cultural trends have led to the reduction of the acreage under rhubarb in West Yorkshire from almost 4000 acres to about 150 acres from which three main growers produce the bulk of British rhubarb. Wilmore (2000) has remarked on that there has the general trend towards cereal production and a consequent decline in the use of wool waste as a fertiliser. A decline in the areal extent of traditional rhubarb fields has thus reduced the extent of habitat treated with wool waste and consequently the opportunities for the growth of wool aliens. CHARACTERISTICS OF THE WOOL ALIEN FLORA OF THE RHUBARB FIELDS OF WEST YORKSHIRE From the comprehensive data presented in Wilmore (2000), it is clear that most of the recording of wool aliens in West Yorkshire in the period 1950-1980 was undertaken on the waste tips of scouring mills in the region. Significantly, all the records prior to 1980 came from localities in ten kilometre grid squares other than SE32 and it was mainly through the fieldwork of John Martin that the flora of the shoddy fields of Ardsley, Rothwell and Oulton in this latter grid square became so well known. His database and herbarium and the publication of his records by Wilmore (2000) are surely one of the major contributions to botanical WOOL ALIEN FLORA OF WEST YORKSHIRE 131 recording in the British Isles in the late- twentieth century and the data allow meaningful interpretations of the nature of the presumed wool alien flora in one of its last refuges. Fieldwork by the author since 2000 has suggested that the special ecology and environment provides a refuge for several groups of uncommon and declining alien plant species, a selection of which is discussed in the following text. Seven species of the Pigweed genus Amaranthus are considered by Wilmore to be especially characteristic of the shoddy fields, whilst formerly being found as grain aliens in docklands and on local authority rubbish tips in the 1950s and 1960s. It is difficult to consider several of them to be wool aliens (A. albus, A. blitoides, A. hybridus and A. retroflexus) for they originate from North America, never a traditional source of imported wool in living memory, but a source of raw cotton, oilseed and grain, commodities that may have been a past vector. Wool imports from South Africa and South America may have resulted in the presence of A. capensis, A. thunbergii and A. quitensis, but the quantities imported have been so small that other vectors such as oilseed or bird seed must be considered as_ strong contenders (Hanson & Mason 1985). One of the rhubarb growers recalls using oilseed waste from the mills of British Oil and Cake Mills Ltd., (BOCM) at Selby in the 1970s (BOCM Pauls 2005). From its early establishment in 1910, the port mill of Selby on the Aire-Calder Navigation, some 30 km to the east, crushed oilseeds to produce vegetable oils for the human food industries and the by-product of this process was oilseed cake, a rich source of protein for feeding to all types of animals. The cake comprised waste from vegetable oilseeds such as soya from the USA and South America, groundnuts from West Africa and cottonseed from East Africa and the Far East, mixed with waste from maize and hard red wheat from the USA. All these provenances could account for the presence of most Amaranthus species recorded as aliens in Britain. But this source for the shoddy field aliens would necessitate either regular replenishment with the waste, for which there is no evidence, or production of viable seed annually for twenty years by A. hybridus to account for many of the later records. This latter situation may be the case for there are fourteen records from 1982 to 1998 passim (Table 1), four from Woodhouse Lane Farm, East Ardsley (SE2924), in which locality I have TABLE 1. COMMONNESS AND RARITY OF AMARANTHUS AND ERODIUM SPECIES IN V.C. 63, POST-1980, IN 10 KM GRID SQUARES SE22 AND SE32 Species Origin Amaranthus hybridus Tropical & North America A. albus North America A. retroflexus Tropical & North America A. blitoides North America A. thunbergii Tropical & South Africa A. viridis Tropical America A. capensis South Africa A. deflexus South America A. quitensis South America Erodium cicutarium Native/Mediterranean E. moschatum Mediterranean E. botrys Mediterranean E. crinitum Australia E. stephanianum China, SE Asia E. brachycarpum North Africa E. chium Mediterranean E. malachoides SW Asia E. ciconium Australia E. cynorum Australia E. laciniatum Mediterranean post 1980 1 km grid Other 10 km Records squares records 14 8 - 3) 3 SE42 3 iy) SE13/60/72 2 2 - 2 2 — 2 2 - 1 1 — 1 l ~ 1 1 = >20 >10 22 SE squares >20 >10 SE11/13 15 8 SE13 8 6 SE13 3 3 - 2 y) = 2 y ~ 2 2 - 1 I ~ 1 1 ~ 1 1 - [Tabulated data derived from an interpretation of cartographic details presented by Lavin & Wilmore (1994) and textual records published by Wilmore (2000)] 132 recorded it in 2002 and 2005. If regular seed set does occur, therefore, one must simply surmise that it is the phospho-nitrophilous nature of the habitat that encourages the growth of such a diversity of species, rather than repeated introductions; that it is ecology which is the determining factor for their ontology. Other species from a variety of families have equally dubious origins. For example, Sisymbrium irio, according to Rich (1991) is “often associated with grain imports and often as a wool alien’, whereas Pearman (2002) considers it to be “formerly a wool alien.” Of the forty-one records cited in Wilmore (2000), twenty-nine are from wool waste dumps and shoddy fields of Yorkshire, and in two localities at East Ardsley (SE2924) and Newton Hill (SE3222), there are respectively ten and nine annual records in the period 1981-94. These data and personal observations in the period 2002-05 indicate that there can be no doubt that this species of Eurasian origin reproduces annually by seed and as such cannot be considered as a wool casual requiring reintroduction from seed in waste wool originating in the Middle East. Yet one must not dismiss a similar origin to the species of Amaranthus, as an oilseed alien maintaining itself in a nutrient-rich, frequently disturbed habitat. The shoddy field habitat could probably not be more contrasting than some of those of the plant’s original London localities, such as the Roman walls and environs of the Tower. A similar diversity of origin is implied for various alien members of the Poaceae, Yorkshire records for approximately 30% (45 species) of which come from this locality. The species are mostly Mediterranean in origin and there is a strong probability that most may be sourced to wool imports, but the rhubarb fields are also the main habitat for Ehrharta longiflora from South Africa, Ceratochloa brevis and C. cathartica from South America, and from tropical regions inter alia, Chloris virgata, Setaria verticillata and Pennisetum clandestinum. Again, the oilseed cake residues from Selby are perhaps more strongly implicated as vectors than wool waste. Certain groups of species with fruiting heads, fruit or seeds that are clearly adapted to animal dispersal by their ornamentation with hooks and spines may be considered with a far greater certainty as wool aliens. Examples of gross morphological adaptations include the species of Centaurea mentioned earlier, the two D. W. SHIMWELL species of thistle, Carduus tenuiflorus and C. pycnocephalus, and Datura stramonium for which the rhubarb fields are probably the main inland locality in Britain. In the Lamiaceae, Marrubium vulgare has hooked calyx teeth, in the Malvaceae, the two commonly encountered species Malva pusilla and M. parviflora, have reticulate and ridged nutlets, whilst most species of Medicago (Fabaceae) have curved or spiraled, spiny fruits. Wilmore (2000) lists eleven species in the genus, of which M. arabica, M. laciniata, M. minima and M. polymorpha are the most common and particularly characteristic of the shoddy fields where they will certainly set viable seed. There are, for example, relatively early records by Kit Rob for M. laciniata from a potato field at Berryhills in v.c. 65 for the period 1958-63, and it was also recorded during the present study in a single rhubarb/cauliflower field at Rothwell (v.c. 63) in the period 2000-05 by the author. Wilmore also records eleven species of Erodium (Geraniaceae) in the Rhubarb Triangle, only four of which were found to be relatively frequent during the past five years’ research (Table 1). All these species are characterized by the beaked or hooked apices of their mericarps, a feature which is no doubt responsible for their dissemination in wool waste. Although the native FE. cicutarium is regarded as native in sandy, coastal habitats, here in the vegetable fields it is certainly introduced and is probably of Mediterranean origin. The same is true of E. moschatum, often the most obvious species, capable of attaining a gregarious growth which makes it the dominant weed in several fields. Both species set good seed regularly but observations on whether E. botrys, of Mediterranean origin, and E. crinitum from Australia, do likewise proved to be inconclusive. FLORA OF THE WASTE DUMPS OF THE UPPER COLNE VALLEY A previously unrecorded use of wool waste in agriculture came to the notice of the author in 2000, namely, its use by several milk, suckling and beef cattle farmers in the Slaithwaite district of the upper Colne Valley (SEO1), where local information indicated that the practise had been in operation for the past twenty years. The origin of the shoddy is the Dewsbury scouring company of Chadwick & Sons and it is used for bedding cattle in stall WOOL ALIEN FLORA OF WEST YORKSHIRE 133 during winter. When the cattle are turned out to grass in late-April, the bedding, comprising a mixture of shoddy, straw, manure and silage is cleared to waste dumps and left to rot. After several years rotting, usually a minimum of five, the mixture is spread on _ pastures previously treated with herbicide and then ploughed in as an organic nitrogen-rich ferti- liser to encourage growth of an Italian ryegrass silage crop. That the environments of the dumps are short-lived is exemplified by the fact that those at Booth Hey Top (locality 2) were spread and ploughed into the surrounding pastures in the winter of 2004. In 2005, only Chenopodium murale and Medicago arabica survived in nutrient rich wall bottoms and a marginal footpath strip, the bank of which yielded a surprisingly luxurious growth of Conopodium majus and Ranunculus bulbosus. The waste heaps are augmented by a new dump of used bedding each year, so that several heaps comprise a sequence of five annual deposits on which a short-term vegetation succession is sometimes apparent. The habitat is extremely nitrogen-rich and Table 2 shows both the nitrogenous nature of wool shoddy relative to other animal by-products and the contributions of the other three components of the habitat. The habitat differs from that of the rhubarb fields of East Ardsley and Rothwell in this respect that the environment is wholly waste dump, lacking adjacent, native arable soil. The other major difference between the two habitats is that the Slaithwaite waste dumps are at least 200 m higher at 245-330 m compared to an altitudinal range of 50-125 m for the rhubarb fields. These two considerations, plus the fact that the waste dumps are simply much smaller in area, accounts for the lower species diversity; no species was present on the dumps and absent in the rhubarb fields, though some such as Chenopodium murale had a seemingly greater frequency of occurrence. A study of the five waste heaps over the period 2001—2005 showed that the components of their flora may be divided into six ecosociological groups: a. a group of common annual weeds typical of most nutrient-rich habitats that occur in abundance over the first three years of the life of the waste dumps: Atriplex patula, Chenopodium album, Epilobium ciliatum, Matricaria discoidea, Polygonum aviculare, Poa annua, _ Persicaria lapathifolia, Sonchus oleraceus and Stellaria media; TABLE 2. PERCENTAGE NITROGEN CONTENT OF COMPOST MATERIALS Source | Source 2 Animal Manures pigeon 4:2 --- fowl 1-6 1-0 dog We) --- duck Ifo} 1-0 pig 0-6 0-4 horse 0-4 0-6 cow 0:3 0-4 Animal By-products feather 15 8 blood meal 15 iD hoof & horn meal 12 12 silk waste 8-4 -- wool shoddy 8 10 fish meal 6:5 9 bone meal 4 4 Vegetable Wastes field bean haulms 1-70 cotton waste egy timothy hay/silage 125 brewing molasses/grain 0-70 potato haulms 0-60 wheat straw 0-50 beet haulms 0-40 Source 1: Primal Seeds (2004). Percentage comp- osition of composts. http://www.primalseeds.org/ npk.htm Source 2: Moor, F (2004). The world of soil: food values of different manures and fertilisers. http:// web.ukonline.co.uk/fred.moor/soil/fertilis/f01.htm b. a group of gregarious perennials that gradually come to dominate after approximately five years: Cirsium arvense, Dactylis glomerata, Elymus repens, Galium aparine, Lolium perenne, Phleum pratense, Rumex obtusifolius, Trifolium repens and Urtica dioica; c. species with helophyte tendencies, both annual and perennial, growing in the moist environment provided by the _ water- absorbent shoddy: Agrostis stolonifera, Alopecurus geniculatus, Impatiens glandulifera, Puccinellia distans and Ranunculus repens; d. cereal species from seed in the straw and cattle feed: Avena fatua, Hordeum vulgare and Triticum vulgare; e€. pioneer species characteristic of high nitrogen environments such as Cheno- podium murale, Datura stramonium and Arctium minus; 134 D. W. SHIMWELL TABLE 3. FREQUENCY OF SELECTED ANNUAL AND BIENNIAL SPECIES OF ECOSOCIOLOGICAL GROUPS E & FON SLAITHWAITE WASTE DUMPS, 2001-05 Species 2001 2002 2003 2004 2005 Records %F C/I Chenopodium murale 1-3,5 1-5 1,2,4,5 1-5 12. Siem all 80 -1-63 Erodium cicutarium 1-5 1,3-5 1,3-5 1,3-5 1,3-5 7\ Vil -0-11 Medicago arabica 1-3,5 1-3 1-3,5 1-3,5 1-5 20 Tah +0-69 Erodium moschatum LP) gs) 1D) od) 55) 13 53 +0-47 Melilotus indicus 1-3 LZ ee 1,3-5 1 12 50 -1:59 Melilotus albus eZ 2 2 29) 1 10 33 -0:20 Geranium molle 153 3 1 13 IES) 8 33 -0-46 Melilotus officinalis 1,4 1,4 13) 4 1 8 38 +0-02 Arcticum minus 4,5 4,5 - 4,5 4 if 28 -0-41 Malva parviflora 1,4,5 - 3,4 3)5) - a 28 nd Malva pusilla - 1,2,4 2 iL - 7 28 nd Geranium pusillum l 1 13) 1 l 6 24 +0:16 Geranium lucidum 1 1 1 l 1 >) 20 +1-42 Marrubium vulgare D 2 yy 12 - 5) 20 -2-02 Medicago polymorpha 1e2 1 - - 4 + 16 -1:34 Datura stramonium - - 2 2 - 3 12 -0:71 Geranium dissectum 1 ] - : 3 1 -0:09 Trifolium arvense I 1 1 - - ) 12 -0-:01 Carduus tenuiflorus i 1 - - - 2 8 -0:14 Erodium botrys - el di 2 2 - 2 8 nd Sisymbrium irio - - 3 - 3 2 8 +0-13 Localities Grid Ref Alt (m) Area (m’) 1. Coalgate, Slaithwaite SE052140 330 3500 2. Booth Hey Top, Slaithwaite SE057137 315 1500 3. Row Farm, Slaithwaite SE063138 285 900 4. Delves Lane, Dowry Farm, Lingards SE077128 265 1500 5. Shroggs Top, Dowry Farm, Lingards SE076129 245 1000 Records = number of records in 2001—2005; %F = % frequency of occurrence; C/I = Change Index as derived from Preston, Pearman & Dines (2002) f. annual and biennial species typical of open habitats, probably introduced as seed from the cleaning of imported wool; divisible into four sub-groups: (i) members of the Fabaceae with nitrogen- fixing root nodules — Medicago arabica, M. polymorpha, Melilotus albus, M. indicus, M. officinalis, Trifolium arvense; (ii) members of the Geraniaceae, including Erodium cicutarium, E. moschatum, E. botrys, Geranium dissectum, G. lucidum, G. molle and G. pusillum; (i111) members of the genus Malva — M. parviflora and M. pusilla. (iv) members of other families — for example, Marrubium vulgare (Lamiaceae) and Carduus tenuiflorus (Asteraceae). “ssp. boraei In terms of the sociological affinities of the six components, Group (a) constitutes a variant of OV33 Polygonum lapathifolium-Poa annua community, the Group (b) assemblage fits into OV25 Urtica dioica-Cirsium arvense community, and Group (c) finds a place in OV28 Agrostis stolonifera-Ranunculus repens community (Rodwell 2000). Groups (e) and (f ) show the closest affinity to OV25 which is seen to contain four of the species in Table 3 (Arctium minus, Geranium dissectum, G. molle and Malva neglecta), plus Amaranthus albus. There are other less significant associations with three species. Specifically, these are OV5 Digitaria ischaemum-Erodium — cicutarium community (with Amaranthus _ retroflexus, Erodium cicutarium and Geranium molle) and OV6 Cerastium glomeratum-Fumaria muralis community (with Erodium WOOL ALIEN FLORA OF WEST YORKSHIRE 135 moschatum, Medicago arabica and Medicago polymorpha). However, the apparent restriction of these two communities to the Bagshot Beds in Surrey and the Scilly Isles and southwest Cornwall respectively, precludes any meaningful insight into sociological affinities. Chenopodium murale is recorded only once in OV30 Bidens tripartita-Polygonum amphibium community typical of the eutrophic margins of lowland ponds and clearly not closely akin to the waste dump habitat in its physiognomy. From the frequency records in Table 3, the waste dumps are clearly a major habitat for this archaeophyte which has a change index of - 1:63 (Akeroyd 2002). It is one of a group of species with a relatively high negative change index that includes Marrubium vulgare (-2:02), Melilotus indicus (-1:59) and Medicago polymorpha (-1-:34). Finally, it is interesting to note that of the seventeen species listed in Group f above, only the three Melilotus species, Erodium botrys, Malva parviflora and M. pusilla are given full records by Wilmore (2000). Nine of the remaining species are relegated to an appendix on account of their Native status in parts of Britain other than Yorkshire, whilst three — Geranium dissectum, G. molle and G. pusillum — are not considered as aliens. These three species of Crane’s-bill are clearly to be considered as wool aliens and probably symbolise the changing face of the woollen manufacturing industry in that most of the home scoured wool, and hence the waste, comes from UK sources. DISCUSSION AND CONCLUSIONS From the foregoing information, it is quite clear that the nature of the UK _ woollen manufacturing industry has undergone major changes in the past thirty years, particularly, in a context relative to the changing flora of Britain, with respect to the change to the predominant use of imported, clean wool and the concomitant decline of British scouring companies. As a direct consequence, the quantities and sources of wool waste available to the agricultural industry have declined dramatically. Further, there is now a predominance of the use of the home clip from Britain and Ireland over imported wool in the few remaining scouring mills, and the shoddy manufacturers are legally bound under EC regulations to use only clean imported rags. All these features have contributed to a major decrease in the size and diversity of the wool alien seed bank entering the country. In addition, the relatively small quantity of wool waste available for use as fertiliser in agriculture has declined drastically and hence, so has its geographical use. Thus, at the start of the twenty-first century, the potential replenishment and continued existence of a wool alien flora is apparently restricted to two localised areas of West Yorkshire. The future prospects for this specialised section of the British flora would thus appear to depend on the continued operation of the two scouring mills and the use of wool waste in the rhubarb growing industry and as in-stall bedding for livestock. In an era of multiple subsidies for many aspects of the processing and production industries there may be a case to be made for the conservation of this traditional industrial and agricultural linkage as a strategy for halting future loss of biodiversity. Perhaps a more practical and immediate contribution to an understanding of the future prospects of this section of the British flora would be the inception of a programme of research into the reproductive capacities of selected species as a contribution to the conservation of biodiversity in the face of changing climate. Four main groups/types would seem worthy of investigation. First, a comparative study of the various species of well represented genera, such as Amaranthus, Erodium and Medicago, should yield valuable data on their relative ability for establishment as stable neophytes. Second, the investigation of plants with high negative change index values, >1-0 (Preston, Pearman & Dines 2002), such as the native Marrubium vulgare (-2:02) and Medicago polymorpha _ (-1:34), the archaeophyte Chenopodium murale (-1:63) and the neophyte Melilotus indicus (-1:59) should provide insights into their conservation. Third, there would seem to be some merit in acquiring information on certain neophytes for which there is a lack of change index data, including Malva parviflora and M. pusilla, Trifolium angustifolium and T. hirtum. Finally, an in- vestigation of the reproductive capacities of native species in this habitat, in comparison with their capacities in other more typical habitats, such as Geranium lucidum, G. molle and G. pusillum and Trifolium arvense, might — form a fourth avenue of research. Apart from generating an important botanical and eco- logical data base, such research would also aid the definition of which species might be termed established ‘wool aliens’ and which considered as mere ‘wool casuals’. 136 D. W. SHIMWELL ACKNOWLEDGMENTS I am grateful to Geoffrey Wilmore for his valuable comments on the manuscript. The following people provided information on various aspects of the production of wool waste and its modern uses: Mark Andrews of Thomas Chadwick & Sons, Eastfield Mills, Dewsbury and Tim Whitaker of Haworth Scouring Co., Cashmere Mills, Bradford on wool scouring and waste; Paul Summers of Cullingworth, Summers & Co. Ltd., Batley and Charles Day of Henry Day & Sons Ltd., Dewsbury on changes in the manufacture of shoddy fibre and textiles; John Hartley, Halifax, shoddy manure merchant; Geoffrey Holmes, David Eastwood and the Oldroyd family of the ‘Rhubarb Triangle’ on rhubarb growing; Brian Bamforth, Row Farm, Tim Garside, Highfield Farm and Brian Shaw, Dowry Farm of Slaithwaite on the use of wool waste in the milk and beef production industry, and for free access to their waste dumps. Finally, Sal, Rosie, Merry and Joe first alerted me to the existence of the waste dumps and provided the stimuli for several subsequent field visits. REFERENCES AKEROYD, J. R. (2002). Chenopodium murale, in Preston, C. D., Pearman, D. A. & Dines, T. D. (2002). New Atlas of the British & Irish Flora. Oxford University Press, Oxford. BEAVER, S. H. (1941). The West Riding of Yorkshire, in STAMP, L. D., ed. The Land of Britain. The Report of - The Land Utilisation Survey of Britain. Part 46. London School of Economics, Geographical Publications Ltd., London. BocM PAULS (2005). A history of British Oil and Cake Mills Ltd. http://www.bocmpauls.co.uk/bocmpauls/ main/page.jhtml (accessed 3 August 2005). CLEMENT, E. J. & FOSTER, M. C. (1994). Alien Plants of the British Isles. Botanical Society of the British Isles, London. Dony, J. G. (1953). The Flora of Bedfordshire. Hitchin Museum, Hitchin Urban District Council. Dony, J. G. (1969). Additional notes on the flora of Bedfordshire. Proceedings of the Botanical Society of the British Isles 7: 523-535. Dony, J. G. (1976). Bedfordshire plant atlas. Borough of Luton Museum and Art Gallery, Luton. Dony, J. G. & Dony, C. M. (1986). Further notes on the flora of Bedfordshire. Watsonia 16: 163-172. FITCHETT, C. E. (1943). Bedfordshire, in STAMP, L. D., ed. The Land of Britain. The Report of The Land Utilisation Survey of Britain. Part 55. London School of Economics, Geographical Publications Ltd., London. HANSON, C. G. & MASON, J. L. (1985). Bird seed aliens in Britain. Watsonia 15: 237-252. HAYWARD, I. M. AND DRUCE, G. C. (1919). The adventive flora of Tweedside. T. Buncle & Co., Arbroath. LAVIN, J. C. & WILMORE, G. T. D. (1994). The West Yorkshire Plant Atlas. City of Bradford Metropolitan Council, Bradford. LEES, F. A. (1888). The Flora of West Yorkshire. Lovell Reeve, London. LEES, F. A. (1941). A Supplement to the Yorkshire Floras. A.. Brown, Hull. LOUSLEY, J. E. (1958). Alien plants introduced by the Yorkshire wool industry. Naturalist (Hull) 1958: 77-81. LOUSLEY, J. E. (1960). How sheep influence the travel of plants. New Scientist 8: 353-355. LOUSLEY, J. E. (1961). A census list of wool aliens found in Britain, 1946-1960. Proceedings of the Botanical Society of the British Isles 4: 221-247. LOUSLEY, J. E. & Dony, J. G. (1952). The travels of plants. Bedfordshire Magazine 3: 185-189. Moor, F. (2004). The world of soil: food values of different manures and fertilisers. http:// homepages. which.net/~fred.moor/soil/fertilis/f0107.htm (accessed 8 November 2004). PEARMAN, D. A. (2002). Sisymbrium irio, in PRESTON, C. D., PEARMAN, D. A. & DINES, T. D. (2002). New Atlas of the British & Irish Flora. Oxford University Press, Oxford. PRESTON, C. D., PEARMAN, D. A. & DINES, T. D. (2002). New Atlas of the British & Irish Flora. Oxford University Press, Oxford. PRIMAL SEEDS (2004). An alphabetical list of random compost materials. http://www.primalseeds.org/npk.htm (accessed 8 November 2004). RICH, T. C. G. (1991). Crucifers of Great Britain and Ireland. B.S.B.1. Handbook No.6. Botanical Society of the British Isles, London. RODWELL, J. S., ed. (2000). British Plant Communities 5: Maritime communities and vegetation of open habitats. Cambridge University Press, Cambridge. SOIL SURVEY OF ENGLAND AND WALES (1977). Soils of South and West Yorkshire, 1:250,000. Ordnance Survey, Southampton. SOIL SURVEY OF ENGLAND AND WALES, (1983). Soils of Northern England, Sheet 1, 1:250,000. Ordnance Survey. Southampton. WOOL ALIEN FLORA OF WEST YORKSHIRE 137 STACE, C. A. (1997). New Flora of the British Isles. Second Edition. Cambridge University Press, Cambridge. WILMORE, G. T. D. (2000). Alien Plants of Yorkshire. Yorkshire Naturalists’ Union, York. WOOL RESEARCH ORGANISATION OF NEW ZEALAND (2005). A history of technological innovation in New Zealand. http://www.techhistory.co.nz (accessed 28 April 2005). (Accepted April2006) a 1 ee Patt at ae : ? i Dn! m - i it i 1 My i 1 uy ey i { 7 f eS | | if I ih I i 3] | | yea I 1 , y Yee | | ¢ r i | 1 te . J i} é rate ’ a , ; ' ‘ ht ran | i Ve i J S i 2 “ : : | (a f UA b { { | , y ‘ j h i | = Je fy } t j {a r | | ‘ . ; hi iN lal , i) | - iii ; ; | ! ft ih | o | r 1} } i | D j ih} : ‘ iN} } | so WV Fe IK , {| , i | own: - i ' | . ail } y . ‘can 5 if 2 x jie ' ; ay | i) 1) \ / Ny 7 a | y igen : Ha MO, ! H Hi uw oon 1} f , ' 7 m1 : ME 3 i i 4 i i iy ; Uy ir, sy | ; iy i : ji al (nae i : Conte ny a) ri 7 1 av 7 ii i 4 i Watsonia 26: 139-144 (2006) 39 Conservation of Britain’s biodiversity: Hieracium riddelsdelli (Asteraceae), Riddelsdell’s Hawkweed T. C. G. RICH* Department of Biodiversity and Systematic Biology, National Museum of Wales, Cardiff CF 10 3NP ABSTRACT Hieracium riddelsdellii Pugsley, Riddelsdell’s Hawkweed, is a rare Welsh endemic plant. It was first found in 1899, was described as a variety in 1907, and raised to a species in 1941. It has been reported from four sites, but is currently only known in two localities in the Brecon Beacons (v.c. 42) and one in Carmarthenshire (v.c. 44), with an estimated total population of about 870 plants. Records for Berwickshire (v.c. 81) refer to another taxon. It qualifies as “Vulnerable’ under the IUCN Threat Criteria. The main threat is the spread of Cotoneaster integrifolius over its Carboniferous Limestone rocky habitats, but all populations would also benefit from relaxation of grazing pressure. It is being cultivated at the National Botanic Garden of Wales and seed has been deposited in the Millennium Seed Bank. KEYWORDS: endemic, IUCN _ Threat lectotype, Wales. Criteria, INTRODUCTION Hieracium riddelsdellii Pugsley (Asteraceae), Riddelsdell’s Hawkweed, is a very rare, endemic plant, which has been reported from localities in the Brecon Beacons (v.c. 42), Carmarthenshire (v.c. 44) and Berwickshire (v.c. 81) (Sell & West 1968; Wigginton 1999). It was reported from four 10-km squares by D. McCosh in the Vascular Plant Red Data Book (Wigginton 1999): these were SN81 Craig-y- nos, SN82 Llyn y Fan Fach, SN92 Craig Cerrig-gleisiad and NT96 Burnmouth (D. McCosh, pers. comm., 2004). Recently, as the status of some of its records and the relation- ship between the Brecon and Berwickshire plants had been revised, further information on its distribution and population sizes was required to inform conservation work. A joint project was therefore set up between the National Museum of Wales, the National Botanic Garden of Wales and the Countryside *e-mail: tim.rich@nmgw.ac.uk Council for Wales to review its current status and determine its need for conservation. In this paper the information is summarised; full details can be found in Rich (2004a, b). TAXONOMY AND IDENTIFICATION Hieracium riddelsdellii was first separated as a distinct taxon by A. Ley who named it as H. ciliatum Almq. var. venosum (Ley 1907). Ley had first found it abundantly on the Craig-y-nos ridge in 1899 and cultivated it thereafter. Later, whilst naming H. repandum Ley (now H. repandulare Druce), he transferred it to that species, retaining it at varietal rank (Ley 1909). Pugsley (1941) raised it to species rank and named it after H. J. Riddelsdell (one of the early collectors), as the name H. venosum L. was already in use for another species. It is a member of Hieracium Section Stelligera Zahn (Section Sub-Oreadea Pugsley). A lectotype for H. ciliatum Almq. var. venosum Ley, for which 4H. riddelsdellii Pugsley is a nom. nov., has been selected by P. D. Sell and C. West as the sheet collected on 1 August 1899 from the high limestone ridge above Craig-y-nos, Breconshire by A. Ley in herb. Ley (currently in CGE) which has Ley’s original description and notes attached; it is hereby designated by P. D. Sell. Some other sheets in BM and BIRM have a different selection noted, which should now be disregarded. SUMMARY OF SYNONYMY: Hieracium riddelsdellii Pugsley, Journal of Botany (London) 79: 181 (1941). = H. ciliatum Alma. var. venosum Ley, Journal of Botany (London) 45: 109 (1907). = H. repandum Ley var. venosum Ley, Journal of Botany (London) 47: 13 (1909). 140 T. C. G. RICH H. repandulare: involucral bracts c. 12-15 mm long, with broad, pale green margins and darker centre, gradually tapering, with numerous stellate hairs especially on margins Involucral bracts c. 10—12(—13) mm long, dark greyish all over, obtuse, with dense stellate hairs on margins and apices G Ligules mostly 20-25 mm — long, hairy at the tips Yi Later rosette leaves ovate- lanceolate or ovate, glandular- denticulate, rounded or subtruncate at base Primordial rosette leaves elliptic, nearly entire FIGURE 1. Ilustration of Hieracium riddelsdellii showing main identification features. A. Whole plant. B— F. Basal rosette leaves. G. Flower. H. Involucral bract. I. Involucral bract of H. repandulare. Scale bars 1 cm. HIERACIUM RIDDELSDELLII 14] TABLE 1. CHARACTERS DISTINGUISHING HIERACIUM RIDDELSDELLIT AND H, REPANDULARE Character H. riddelsdellii Base of rosette leaves Involucral bracts (Figure 1, H) Capitula Key taxonomic features of H. riddelsdellii are the oval basal rosette leaves which are glandular-denticulate with a rounded or sub- truncate base and sometimes large teeth, the rather slender usually leafless stem, the medium-sized heads which are rounded below, the dark green, mostly obtuse/subacute involucral bracts with dense stellate hairs on margins and apices (the latter like a white tuft in fresh material), the hairy tips to the ligules, and the yellow styles (Figure 1). Sell & West (1968) noted that H. riddels- dellii is very similar to H. repandulare and could have been derived from the same parent, and that there are no very close allies of either. The differences between the two species are set out in Table 1; the best characters are the size, Shape and hairiness of the involucral bracts. Pugsley (1948) suggested H. riddelsdellii could also be distinguished from H. repandulare in lacking the pronounced out-curved teeth at the base of the rosette leaves and the more compact inflorescence. However, during field work it was found that the leaf bases of both species are significantly more variable than noted by Pugsley, and toothing of the leaves is unreliable for identifying them. The differences in com- pactness of the inflorescence are also more visible in cultivated plants than in the field. Sell & West (1968) noted that plants at Burnmouth, Berwickshire (v.c. 81) matched the Brecon plants exactly morphologically and could be regarded as conspecific, though perhaps of different origin. More recently, this diagnosis has been questioned (Braithwaite & Long 1990). The original record was based on a specimen in CGE which is no longer in the H. riddelsdellii folder and has been re- determined, though P. D. Sell cannot currently recall to which species (pers. comm. 2005). D. McCosh has failed to refind H. riddelsdellii- Rounded to truncate, entire or with a few small teeth, younger rosette leaves sometimes with more pronounced teeth, usually green or partially flushed red c. 10-12(—13) mm long, greyish over most of involucral bract, obtuse-subacute, with dense stellate hairs on margins and apices Smaller, ligules up to 28 mm long H. repandulare Rounded to truncate (rarely cordate), entire or toothed with small or pronounced teeth, often glaucous, sometimes green c. 12-15 mm long, with broad pale green margins and darker centre, gradually tapering, with numerous stellate hairs especially on margins (Figure 1, I) Larger, ligules up to 34 mm long like plants on several visits to the Burnmouth populations; H. britanniciforme Pugsley is the most common plant at Burnmouth, and another nearby locality has a Hieracium species with noticeably pilose ligules but which is not H. riddelsdellii. DISTRIBUTION The historical records traced from the literature and herbaria (BM, BIRM, CGE, E, LIV, MANCH, OXF and NMW) are summarised in Table 2. It has been confirmed in three sites, and recorded doubtfully in another. Ley (1907, 1909) reported it only at Craig-y-nos; he had also collected it at Llyn y Fan Fach but included it under H. repandulare. There are also. many duplicates of his cultivated specimens from Craig-y-nos 1903-1910. Records for Craig Cerrig-gleisiad are errors for H. repandulare. The historic records were used to direct the field surveys. In addition, other suitable areas were searched in the Upper Tawe valley. CRAIG-Y-NOS RIDGE, V.C. 42 BRECON (SN8314 AREA) Craig-y-nos Ridge (also known as Cribarth) is a large Carboniferous Limestone hill on the west side the Tawe Valley, with low broken cliffs and screes along the south-east side and quarries on the top and at the east end of the hill. Hieracium riddelsdellii was found scattered and sometimes abundant throughout much of the site, especially on the unquarried rocks. In 2004, 388 plants in flower were counted, but there were many vegetative rosettes suggesting a total population of about double that, c. 750 plants (Rich 2004b). CRAIG RHIWARTH V.C. 42 BRECON (SN8314 AREA) There is a specimen from Craig Rhiwarth, 8 July 1906, A. Ley in CGE, but I am T. C. G. RICH 142 AIT Wa MIWN ‘HONVW ‘ATT “A990 ‘Wald ‘WA MIN ‘AXO ‘ATT AD) WE Wa MIN ‘AXO ‘ATT (244101991) ADO ‘INGA ST cOO8NS ‘A][NS uleul ‘SuI[poos | pue sjueyd 6 ‘yoy ue A UAT VICOO8NS 49U409 UJOISOM UT AT[NS UTeUL ‘AAOge sJJI[O ‘yoRy uey A UATT dIysuoyyeULeZ ‘ueyse, ue UAT] A SIeID I9L9S8NS Jvou Arsenb posnsip ‘jy [euag CIS pur pIps ‘pIE8NS Auenb pur ospu sou-A-s1esD OSTIP8NS SJJt[9 OuOsouUT] “soU-A-STeID wey jeM-A-jueg saoge Arenb pio sou-A-3Ieia Aisenb sou-A-31elg OSTIP8NS ‘Aq Asrenb ‘apyyseg sou-A-31eIg Aisenb pjo ‘sou-A-31eig sou-A-3Iely sou-A-3Iely asp sou-A-3IeID Spl soU-A-3IvID dSplLI SOU-A-3IvID Ajjuepunqge “VW OSTI ‘JO ISOM ISPLI DUOJSOUNT] YSty ‘sou-A-sIeID sojou pure 901n0S aS NWTIAGSTAAd1a WOIDVAAIH JO SAXOOda WOLAV das ¢ ATaV yor “DOL 7OOT/9/LTZ SOTA V A 0961/L/07 Kay Vv 906I/L/P (pp 9°A) yory ue A UAT] SITTIN 'N ‘f S96I/8/ET (Tp (9°A) WIAMUog pueH ‘O'S WM 'D'O'L 0007Z/9/S I SITTIN “Nf €961/9/I SOTA “V “A 8C61/9/SI sUIUMOIG “YA LS6I/LITT sMoIpuy “A VO CS6I/S/6Z 19S ‘d‘d ES6I/9/ET usAey “Ff ES6I/9O/E KOT ‘VW 906I1/L/8 uojury “4 “7 906T/LIE [PPSTOPPra ‘fH 2 AOT V +061/9/TI Kay Vv 7061/9/01 Kay ‘VW 0061/L/9 Ko] “VW 6681/8/1 (Zp OA) YeQLIig/sou-A-sIeID 10} [OD aep/Ayeo0'T HIERACIUM RIDDELSDELLII unconvinced of its identity. Craig Rhiwarth is a very likely locality being directly up-wind and on the opposite side of the valley to the Craig- y-nos Ridge. The site has been searched three times since 2000, and no material has been found. PENWYLLT, V.C. 42 BRECON (SN8516 AREA) There are numerous disused Carboniferous Limestone quarries around PenwyJllt, all but one of which with no access have been searched. The Twyn Disgwylfa Quarry (SN856161) has been markedly enlarged since H. riddelsdellii was recorded by J. N. Mills in 1965 (Table 2), and no plants could be found in 2004. Hieracium riddelsdellii was however found in the two adjacent disused quarries to the south, with 105 plants flowering in the larger quarry at SN856156, and four plants flowering on west-facing cliffs in the smaller quarry at SN856157. LLYN Y FAN FACH, V.C. 44 CARMARTHEN (SN800215) The Old Red Sandstone crags above Llyn y Fan Fach form a cirque which has had at least fourteen species of Hieracium reported, though only about half this number have been collected in recent years. On 27 June 2002, nine plants and one seedling of H. riddelsdellii were refound in three close groups on the low cliffs at the bottom on the main gully above the western end of Llyn y Fan Fach, where H. riddelsdellii had previously been localized by B. A. Miles. Hieracium riddelsdellii is thus currently known from three populations in two 10-km Squares in two vice-counties, with a total estimated population of 870 plants. One pop- ulation at Twyn Disgwylfa Quarry has been destroyed. A 10-km square distribution map is given in Figure 2. LIFE CYCLE AND ECOLOGY Like most British Hieracium species, H. riddelsdellii is a polycarpic perennial which reproduces by seed. The main flowering period is mid May to mid June, and seeds ripen about 3—4 weeks after flowering. It is probably an obligate apomict: the stamens and styles were excised on one of the two flowering heads in bud on the one plant available in cultivation in 2004, which did not prevent seed set (further larger scale trials are required). The mean potential pollen viability assessed using Alexander’s Stain (Alexander 1969) was 57:2% =eowoomsse) (ni — 25) (H. Cleall & Tf. Rich, 143 peal | 33 FIGURE 2. National distribution map of JH. riddelsdellii, plotted using DMAPW by Alan Morton. @ 2002-2004. x = error. unpublished). The seeds have a small pappus and are wind-dispersed. DNA analysis of five H. riddelsdellii plants using amplified fragment length polymorphism (AFLP) during an in- vestigation of genetic variation in Hieracium cyathis (Ley) W. R. Linton showed each plant was genetically different (M. D. Lledo, pers. comm. 2006). The associated vegetation was recorded in seven representative 2 m X 2 m quadrats. The vegetation was usually open (mean percentage cover 9%, range 5—20%), short (mean height 5 cm, range 2-10 cm), and on steep slopes (mean slope 79°, range 70-90°) of all aspects. In terms of British Plant Communities (Rodwell et al. 1991-2000), the vegetation was generally the OV39 Asplenium trichomanes — Asplenium ruta-muraria community and the CG10 Festuca ovina — Agrostis capillaris — Thymus praecox grassland. Soil pH was measured with a pHep2 Hanna pocket-sized pH meter in a 50:50 mixture with distilled water on soil samples collected from around the roots. Five pHs of soils measured from the limestones of Craig-y-nos area were 695769) 7-059 7-1) and: 7-7, and” pH 6:5) was measured from soils derived from the Old Red Sandstones at Llyn y Fan Fach; these indicate H. riddelsdellii is a calcicole. | In 2000, 2003 and 2004, many flowering heads were infected by a small grub causing distorted growth and sterility (Pugsley 1948 noted this for Hieracia generally), and in 2004 many plants were heavily infested with aphids which were unusually abundant that year. 144 MiCcsG2RICEH CONSERVATION Under the IUCN (2001) Threat Criteria, ZH. riddelsdellii qualifies as ‘Vulnerable’ (total population less than 1000 individuals, highly restricted area of distribution). It is not protected under Schedule 8 of the Wildlife and Countryside Act 1981, but is included in the Vascular Plant Red Data Book (Wigginton 1999). All the H. riddelsdellii sites are in the Brecon Beacons National Park, which gives them a limited degree of protection. The population at Llyn y Fan Fach is within the Mynydd Du (Black Mountain) S.S.S.I.. The Penwyllt population is within the boundary of the Ogof Ffynnon Ddu S.S.S.I. which is designated for its cave system but not for its surface biological interest. The main population at Craig-y-nos is not within an S.S.S.I., and is clearly a priority for designation to conserve H. riddelsdellii as it contains 86% of the population. All sites except for the northern of the two quarries at Penwyllt are heavily sheep-grazed, and H. riddelsdellii is largely confined to rocks out of the reach of sheep (most Hieracia are sensitive to grazing, and H. riddelsdellii is no exception). As a plant of open rocks and cliffs, generally no management is required provided these habitats stay free from scrub and are ungrazed. If grazing is relaxed significantly, it could spread onto adjacent rocky grasslands (all Hieracia present would probably benefit from this). The only immediate threat to the populations is the continued spread of the alien Cotoneaster integrifolius in the quarries at Penwyllt which already requires control. There are no immediate threats to survival of H. riddelsdellii at either Craig-y-nos or Llyn y Fan Fach, though longer term intensive grazing will limit the ability of the population to spread. Two collections of plants grown from seeds are currently being cultivated at the National Botanic Garden of Wales, and seed from 39 plants has been deposited in the Millennium Seed Bank. ACKNOWLEDGMENTS This work was jointly funded by the Countryside Council for Wales, the National Botanic Garden of Wales and the National — Museum of Wales as part of a collaborative project into conservation and ecology of critical species. I would like to thank the Keepers for access to the herbaria and libraries, Wendy Atkinson, Scott Hand, Richard Lester, Lola Lled6, Lindsey Loughtman, Serena Marner, Douglas McKean, Gina Murrell, Peter Sell, Charlie Stirton, Sarah Whild and Leander Wolsten- holme for their help, Alan Orange for identifying bryophytes and lichens, Helen Cleal for data on pollen viability, and David McCosh for naming plants and supplying information from the Hieracium database. REFERENCES ALEXANDER, M. P. (1969). Differential staining of aborted and non-aborted pollen. Stain Technology 44: 117— 1228 BRAITHWAITE, M. E. & LONG, D. G. (1990). The botanist in Berwickshire. The Berwickshire Naturalists Club, Berwick. IUCN (2001). IUCN Red list categories and criteria. Version 3.1. YUCN, Gland. LEY, A. (1907). Hieracium notes. Journal of Botany 45: 108-112. Ley, A. (1909). Brecon and West Yorkshire hawkweeds. Journal of Botany 47: 8-16 and 47-55. PUGSLEY, H. W. (1941). New species of Hieracium in Britain. Journal of Botany 79: 177-183 and 193-197. PUGSLEY, H. W. (1948). A prodromus of the British Hieracia. Journal of the Linnean Society of London (Botany) 54: 1-356. RICH, T. C. G. (2004a). Distribution and conservation of Hieracium riddelsdellii, Riddelsdell’s Hawkweed. Unpublished interim report to Countryside Council for Wales. RicH, T. C. G. (2004b). Distribution and conservation of Hieracium riddelsdellii, Riddelsdell’s Hawkweed: additional information 2004. Unpublished report to Countryside Council for Wales. RODWELL, J. S. et al., eds. (1991-2000). British plant communities. Volumes 1-5. Cambridge University Press, Cambridge. SELL, P. D. & WEST, C. (1968). Hieracium L., in PERRING, F. H. ed., Critical Supplement to the Atlas of the British flora. B.S.B.1., London. WIGGINTON, M. J., ed. (1999). British Red Data Books. 1. Vascular Plants. 3rd ed. Joint Nature Conservation Committee, Peterborough. (Accepted February 2006) Watsonia 26: 145-169 (2006) 145 The status and conservation of stoneworts (Characeae) in West Gloucestershire (v.c. 34) and North Somerset (v.c. 6) R. V. LANSDOWN?*! 45 The Bridle, Stroud, Gloucestershire GL5 4SQ N. F. STEWART*” Cholwell Cottage, Posbury, nr. Crediton, Devon EX17 3QE C. KITCHEN and M. A. R. KITCHEN The Cottage, Bevington, nr. Berkeley, Gloucestershire GL13 9RB ABSTRACT West Gloucestershire and North Somerset are nationally important for stoneworts and although there are diverse assemblages at some sites, the area is particularly important for the strong populations of several rare species. The most important areas are flooded gravel pits in the River Severn floodplain at Frampton-on-Severn, a complex of dew-ponds and pits created through extraction of strontium in the Wickwar — Yate area and ditches in grazing marshes in the Gordano Valley. KEYWORDS: Chara, Nitella, Nitellopsis, Tolypella, distribution, conservation. INTRODUCTION North Somerset (v.c. 6) and West Gloucester- shire (v.c. 34) (referred to hereafter as the region) include a number of sites supporting diverse or large populations of stoneworts (charophytes). These sites are mainly very small and discrete and, as discussed further below, the remainder of the region has little in the way of suitable stonewort habitat. Survey and literature reviews by J. A. Moore and R. H. Bailey in the mid-1970s (Moore & Bailey 1986) and by N. F. Stewart (N.F.S.) and R. V. Lansdown (R.V.L.) since the early 1990s have shown that, although some important sites and Species appear to have been lost, many of the more important sites and the most vulnerable species still occur and the region can be considered to be of national importance for stoneworts (Stewart 2004). *l6_mail: rlansdown@ardeola.demon.co.uk *76-mail: nfstewart @freeuk.com This article presents an account of the status and distribution of charophytes in the region, supported by a comprehensive list of records. The information presented here is_ based on compilation of data from the literature, herbarium specimens, the database of the Biological Records Centre, material submitted to N.FE.S. for determination, the databases of the Bristol Naturalists Society and Gloucester- shire Naturalists Society records held by the M.A.R. and C.K. as well as personal records of all the authors. CHAROPHYTE HABITAT The region comprises the Forest of Dean, the lower, tidal part of the River Severn and its historic floodplain, the southern Cotswolds and the coastal plains south through the Mendips and the Somerset Levels to Bridgwater Bay. It overlaps with all or part of the current ad- ministrative counties of Somerset, North Somerset, Bath and North-east Somerset, Bristol, South Gloucestershire and Gloucester- shire. The landscape is dominated by the hills of the Cotswolds, the Mendips and the Poldens to the east of the River Severn, the Severn Estuary and the Forest of Dean to the west. Much of the region involves the low-lying areas of the Severn Vale and the coastal plain, characterised by pasture and complex field systems with small copses. The hills are broken by a small number of larger rivers, such as the Parrett, the Brue and the Axe in the south and the Little Avon in the north. The former floodplains of the larger rivers have created 146 R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN much of the suitable wetland habitat for charophytes, particularly in the Somerset Levels. Geologically, the region is one of the most varied districts of Britain, for with the exception of the Ordovician, Cretaceous and possibly the Permian, there are surface exposures of every geological formation from the Cambrian to the Jurassic (Kellaway & Welch 1948). The underlying geology has a very strong influence on the distribution of charophytes. One of the reasons for its importance for charophytes is that most of the region is predominantly calcareous, characterised by the inferior oolite of the Cotswolds to the carboniferous limestone of the Mendips, with calcareous clays in the vale of the River Severn (Green et al. 1997; Green et al. 2000). However, there are also important outcrops of acid strata, in particular the coal measures of the Forest of Dean, with an outlier forming the Bristol Coalfields. Overlying the hard geology, most of the coastal plains are dominated by deep alluvial clays, with occasional surface peat deposits in areas such as the Gordano Valley and parts of the Somerset Levels. Within the region, areas with wetland habitats suitable for charophytes tend to be fairly discrete and mainly restricted to the lowlands of the Severn Vale and the coastal plains. There are only a few records from the Forest of Dean, the Cotswolds, the Mendips and Poldens, most of which are from anthropogenic habitats, i.e. habitats that have developed as a result of human influence. The main habitats supporting charophytes in the region are: e Pools remaining from coal and metal ore extraction in the Forest of Dean. e Gravel pits and some ditch systems in the Severn Vale north of Sharpness, particularly in the area from Over to Walham and around Frampton-on-Severn. e Canals, artificial lakes and a few farm ponds on the plateau of the Cotswolds. e Farm ponds and strontium pits in the Wickwar — Yate area. e Predominantly artificial water bodies, such as canals and ornamental lakes, in and around urban areas associated with Bristol and Bath. e The wet pasture and ditches (rhynes or rhines) in the Gordano Valley. e Large artificial lakes such as Chew Valley Lake and Blagdon Lake. e Ditches serving as “wet fences”, scattered standing water bodies and canals in the Somerset Levels, particularly in areas such as Southlake, Butt, Kenn and Nailsea Moors. e Dune slacks on the coast between Weston- Super-Mare and Bridgwater. The more important sites are described in detail below. IMPORTANT STONEWORT AREAS NORTH SOMERSET (V.C. 6) The Somerset Levels The Somerset levels were formed in a large inlet as the sea receded, allowing development of peat and fen woodland (Green et al. 1997). Originally, they would have comprised a vast complex of wetland habitats with smaller dry areas on higher ground. With drainage and cultivation, the wetland habitats have been reduced and are now mainly represented by ditches serving as “wet fences”, with isolated wet grassland, pools and fen. Whilst the ditch systems generally do support some important charophyte populations, the most important sites are those where the ditches are associated with wet pasture. In most recent surveys of the aquatic plants of the ditches in the levels, charophytes have not been identified to species level (often being recorded simply as “Chara sp.) and there is a need for more informed survey. There are scattered records for C. contraria, C..globularis, C. hispida, C. virgata, C. vulgaris, Nitella mucronata and N. opaca as well as old records for WN. translucens, Tolypella glomerata and T. intricata. Records are generally rather scattered, with concentrations in the King’s Sedgemoor area, Kenn Moor and the Nailsea and Tickenham Moor complex. Two sites currently support important populations of T. prolifera: South- lake Moor is an area of wet grassland and grazing marsh on alluvial clays with some peat, where 7. prolifera has been recorded from around 14 ditch sections. Butt Moor is another clay moor with areas grazing marsh and wet grassland with T. prolifera in a number of ditch sections. The Gordano Valley The Gordano Valley extends from Avonmouth, south west to just north of Clevedon. It is mainly an area of grazing marsh defined to the north-west and south-east by steep limestone CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET 147 hills which feed strongly calcareous water into the system. There are charophyte records from four main areas within the valley; ditches in the Portbury area (near Portishead) and three of the moors within what was one continuous wetland complex; Clapton Moor, Weston Moor and Walton Moor. The northern half of the valley has mainly been converted to agriculture and much of Weston Moor has been lost under a refuse tip. However, part of the valley floor is a National Nature Reserve and includes three Avon Wildlife Trust Reserves as well as much being designated as SSSI (Green ef al. 2000). The arterial drains are nutrient enriched but many lesser ditches have a good aquatic flora, particularly Walton Moor and the remnants of Weston Moor (Stewart 2004). The charophyte horas pesull the tichest in’ the region. Chara hispida, C. vulgaris var. papillata and Tolypella glomerata were recorded from Portbury between 1887 and 1922 but there are no recent records of charophytes from this area; Both Walton and Weston Moors still support Chara aculeolata, C. globularis, C. hispida, C. virgata, C. vulgaris and N. flexilis agg. but there has been no record of T. intricata since 1989. Clapton Moor still supports C. virgata and C. vulgaris var. papillata but appears to have lost C. contraria, C. globularis, C. hispida and Tolypella glomerata. In the species accounts, records for all sites within the Gordano Valley have been included under one heading, subdivided only where specific site details are given. WEST GLOUCESTERSHIRE (V.C. 34) The Wickwar — Yate Area This area is situated on the edge of the Severn Floodplain, toward the south of v.c. 34. It contains a much higher than average (Williams et al. 1998) number of ponds (over 450 in the 10 km square ST78) and consequently has provided a high proportion of the records of charophytes for v.c. 34. The ponds derive from strontium sulphate extraction, cattle watering and a small amount of quarrying and mining. The area from which the records come can best be defined by a line running from Rangeworthy in the west, through Wickwar to the base of the Cotswold escarpment below Hillesley, then following the base of the escarpment south through Horton to Old Sodbury, west through Chipping Sodbury to Yate and back to Rangeworthy. In the species accounts a number of named sites shown on the Ordnance Survey 1:25,000 Explorer map (No. 167) have been included within this broad area; these include farms such as Cherryrock Farm, Goose Green (although the farm has been lost to dev- elopment), Hall End Farm and Leech Pool Farm, as well as other features such as the village of Little Sodbury End, Vinney’s Lane (running north-westward from Horton), Yate Court and Yate Rocks. The core of the area is characterised by a complex of commons, including; Assley, Hawkesbury, Inglestone, Kingrove and Sodbury Commons. The commons border Lower Woods SSSI/NR which includes the following named sites referred to in charophyte records: Green Trench, Horton Great Trench, Lance Coppice, Lower Woods Lodge, Spoil Coppice and Stonybridge Wood. Lower Woods NR lies mainly on the Lower Lias clays, with exposures of Keuper and Tea Green Marls, It has a long history of human use, including a roman villa nearby just south of Vinney’s Lane, as well as historic active management. This historical use has resulted in frequent flooded relict features, such as saw-pits, whilst Horton Great Trench may have been a roman road, was probably a drove road and was still a major highway until at least the 1700s (Gloucester Wildlife Trust 1997)» Celestine (strontium sulphate) occurred in irregular masses in the Keuper Marl; it was largely worked by hand and formerly sent to Germany for use in refining sugar from sugar- beet (Kellaway & Welch 1948). It was also used to give red colours to pyrotechnics, but is now mainly used in ferrite for radios and in glass for TV tubes (A. R. Lansdown pers. comm.). The pits resulting from this work were generally small and fairly deep (often over 2 m deep). Consequently, those which have not been actively infilled for agriculture or development have not been particularly affected by succession and most still hold water. They extend from Little Sodbury, Horton, Wickwar and Yate, to the Inglestone Common area. In addition to these ponds, there are a very large number of dew-ponds mainly located at the junctions of two to three fields, particularly on the margins of the commons. A survey of 368 ponds in ST78 between 1996 and 1999 showed that 127 (35%) had been infilled and 36 (10%) had succeeded to terrestrial habitats. Of the remainder, 84 (23%) were under more than 50% shade and only 17 (5%) were sufficiently poached to provide suitable habitat for charophytes such as Tolypella intricata (Stewart & Lansdown 1999b). During the early 1900s, many of the pits were surveyed by people such as J. W. White, 148 R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN I. M. Roper and C. I. and N. Y. Sandwith and most of the pre-1994 records derive from this period. However, few records give sufficiently detailed site descriptions for the individual pit to be matched to current water bodies and so it is difficult to assess whether individual populations still occur. The available data would suggest that Chara vulgaris (including all three common varieties) and C. virgata are as abundant now as they ever have been. Detailed surveys for T. intricata mean that although populations vary between years, it appears to be as abundant now as it ever has been. The type locality for the var. gracillima of Nitella mucronata is in this area, in a strontium pit near Yate Court. The pit appears to have been infilled for development and the plant has not been recorded in the area since the early 1900s. Two other taxa appear to have been lost from the area; both C. globularis and N. opaca were recorded from a number of sites around Yate in the early part of the twentieth century. The area around Yate extends over part of the Bristol coalfield and formerly supported taxa more typical of acid habitats, such as Apium inundatum and_ Baldellia ranunculoides (Green et al. 2000). Unfortu- nately most of the common land and associated wetlands in this area, such as Yate (or Westerleigh) Common, were lost to agriculture during the second World War (Green ef al. 2000), or to the increasing sprawl of dev- elopment around Yate and Chipping Sodbury. Frampton Pools SO7407—7507 Frampton Pools SSSI is a complex of three large and a number of smaller shallow lakes created by gravel extraction, located approximately | km from the flood bank of the Severn Estuary. The lake complex was designated an SSSI in 1974 to protect wintering wildfowl and the developing flora. Many of the lakes are fringed by young woodland, which shades the margins, however in places gravel beaches occur leading down from short, sheep- grazed pasture. Nitellopsis obtusa occurs in a broad band running parallel to a long gravel beach on the north-western side of a lake with a surface area of approximately 21 ha, which is at least 2 m deep in parts. Associated species included Ceratophyllum demersum, Elodea canadensis, and E. nuttallii. Other aquatic plants in the lake included Chara globularis, C. vulgaris, Potamogeton pectinatus, P. pusillus and Ranunculus circinatus. Mendip Reservoirs The importance of the Mendip Reservoirs has only recently come to light as a result of surveys by University College, London in 2003. Of particular note was the discovery of strong populations of WNitellopsis obtusa in both Blagdon Lake and Chew Valley Lake. Undoubtedly there is a need for further exploration of these and other reservoirs in the Mendips. The underlying Carboniferous Limestone results in strongly calcareous water which favours the stoneworts and helps to reduce the effects of nutrient enrichment washed in from the surrounding farmland, although this is still a concern. Stoneworts so far recorded are N. obtusa, Chara contraria, C. globularis and C. vulgaris in both Blagdon Lake and Chew Valley Lake, with C. contraria and C. globularis in Cheddar Reservoir. CHAROPHY TE CONSERVATION Stewart (2004) lists six sites within the region as being important for stoneworts. Of these, the Somerset Levels are considered to be of European importance, Frampton Pools, the Gordano Valley, the Wickwar — Yate area (as Inglestone Common area) and Richard’s Wood at Over are considered to be of national importance and the Thames and Severn Canal is of local importance. The Mendip reservoirs would also have qualified for inclusion but information was not available at the time of publication. Parts of all these sites, except the Thames and Severn Canal are under some level of protection for various reasons and stone- worts are taken into account in _ their management. The Thames and Severn Canal has been out of use since the middle of the twentieth century and is currently succeeding to terrestrial vegetation. However, there are plans to restore parts of the canal and eventually to restore the entire length to navigation. This would comprise the Stroudwater and Thames and Severn Canals, linking the Gloucester and Sharpness Canal at Saul Junction to the Thames at Lechlade. Currently, the future for charophytes in the canal is bleak; but restoration and low-intensity, or no boat use would be extremely beneficial and could restore good populations of all known taxa. Experience from other canal systems, such as the Kennet and Avon Canal, suggests that CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET 149 restoration to anything more than light boat traffic is likely to lead to a more or less total loss of submerged aquatic vegetation, including charophytes. The only conservation action taken specifically for charophytes in the region has involved survey and low level monitoring of Tolypella intricata and T. prolifera. This has led to the development of a _ reasonable understanding of the ecology of the species and proposals for management, although few of these actions have been implemented to-date (Stewart & Lansdown 1999a and b, Williams & Stewart 2002, Williams et al. 2003). Unfortunately, the management plan for Lower Woods (Gloucestershire Wildlife Trust 1997) was produced shortly before detailed information on T. intricata became widely available and this species is not mentioned. Subsequently, the Gloucestershire Wildlife Trust has been involved in the development of conservation management plans for the species. Charophyte taxa such as C. virgata, C. vulgaris and probably C. contraria have the ability to colonise new habitat as it becomes available, whilst at the same time forming a less important part of the flora of established wetland systems. Although some populations of these species in the region have been lost, they are still frequent and in some areas are probably increasing. By contrast, taxa such as C. aculeolata, C. globularis and C. hispida tend only to occur in permanent water bodies. These species are now very local within the region and are vulnerable because of the limited number of small populations. Nitella translucens appears to be extinct in the region, as does Tolypella glomerata although there are some areas, such as the ditches near Portbury and water bodies on the New Grounds at Slimbridge which could hold populations of the latter and merit survey. Nitella opaca is now restricted to a few small populations, however it is likely that this species has been under- recorded in places such as the Forest of Dean and the Somerset Levels. The impression gained is that some species have undergone significant declines since the early part of the twentieth century. However, this impression must be put in the context of the limited number of surveys carried out by botanists capable of identifying charophytes in the field. Only the Wickwar — Yate area, Southlake Moor, Butt Moor and the Gordano Valley have been subject to intensive survey involving field identification. Records of “Chara sp.” from a variety of sites throughout the region would suggest that more thorough survey could re-discover a number of popu- lations of stoneworts, while recent finds of species such as Tolypella prolifera and Nitellopsis obtusa would suggest that some of the re-discoveries are likely to involve species that are of conservation concern. As a minimum, the steps needed to ensure the conservation of existing populations of charophytes in the region involve regular monitoring surveys of suitable habitat in all sites of European and national importance (sensu Stewart 2004). The results of these surveys should be employed to inform decisions on whether or not to intervene with active management for charophytes. In addition, the information gained from surveys should be published and compiled to contribute to the information base on the ecology of these species. Any further development in the Wickwar — Yate area should be subject to environmental impact assessment, giving particular consideration to reducing the loss of waterbodies and remnant common land. The restoration of the Thames and Severn Canal and works undertaken on the Hereford and Gloucester Canal (referred to in the older literature as the Newent Canal) need to be monitored, if nothing else, to record the response of aquatic plants. SPECIES ACCOUNTS Apart from the obvious benefits that derive from compilation of botanical data such as enabling assessment of changes in _ the distribution or status of species and informing conservation action, this article has been prepared to serve-two main aims: to enable an assessment of the current status of stoneworts in the two vice counties and to enable the monitoring of trends in individual populations. To some extent, these two aims dictate the format of data presentation. Rather than simple lists of sites with the date and source(s) of the records, we have reproduced as much detail as was available in each original record to increase the chance of relocating individual populations. Thus, for example, although there is doubt about their validity, we have decided to make it clear where material has been identified to varietal level. In the case of C. vulgaris to list the named variety against each record was impractical and risked doubling the length of this article we have therefore 150 R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN separated out all records identified to variety consistently throughout the article. Records are presented under the species or varietal name following the taxonomy of Bryant et al. (2002). To facilitate assessment of the current status of each taxon, sites for which there are records during the last ten years (..e. since 1994) are listed separately. Where there are post-1994 records for sites where the taxon was recorded before 1994, all records are listed as post 1994. For the same reason, records from the same named site or grid reference are grouped. All material was determined by the recorder unless otherwise stated. The records are arranged as follows: e The ten-kilometre square. e The 6-figure National Grid Reference, where known. e The location as given by the recorder. e The date of the record (where available) and any other useful details. e The initials of the finder e.g. C.LS. A full list of initials and the names to which they refer is provided in the appendix. e [det.] indicates that a specimen was identified by someone other than the finder. e ! indicates that a specimen has been seen by N. F. Stewart and the identification confirmed according to current taxonomy. e Location of any herbarium specimens (in bold), where relevant. e Published sources of the record. Records are listed in chronological order within sites and by the grid reference where more than one record was made in a single year. Any notes added by the authors and in particular, where the site is known but the grid reference was not given by the original recorder, these are given in square bracket []. (BRC) indicates that the only source that we have found for the record is on the Biological Records Centre database at the Centre for Ecology and Hydrology, Monk’s Wood. Synonyms have been included only when they have been used in the two vice counties covered here and do not represent a comprehensive list, for which see Bryant et al. (2002). CHARA L. Chara aculeolata Kitz. Hedgehog stonewort Synonyms: C. pedunculata Kiitz.; C. hispida L. forma polyacantha (A. Braun) R. D. Wood. In Britain, C. aculeolata is nationally scarce and restricted to a few highly calcareous sites, mainly in England. Within the region it has only been confirmed from Walton and Weston Moors, in the Gordano Valley. Material from Nailsea Moor (only a few kilometres south of Walton Moor) in 1880 was recorded as C. hispida var. polyacantha, which probably refers to this species; unfortunately it has not been possible to confirm this record. Populations in the Gordano Valley are re- stricted but appear to be fairly healthy. It may be dependent upon regular ditch or pool maintenance and must be _ considered vulnerable. V.C. 6 BEFORE 1994 ST47 [-Nailsea Moor (as C. hispida var. polyacantha), 1880, A.Le., E.H.R. [det. J.W.W.] (White 1912; Marshall 1914)]. 1994 ONWARDS ST47 - Gordano Valley: - Walton Moor: 10 September 1903, J.W.W. [det. J.G.] (BM!, MANCH!, NMW!, RNG!, SLBI!) (Willis & Jefferies 1959); peat ditch, 10 September 1904, J.W.W. [conf. A.B.] [conf. H. & J.G.] (BM!, MANCH!, NMW!, SLBI!) (Groves 1905; Willis 1989); ST435727 ditch No. 54, 20 October 2002, N.E.S. et al. - Weston Moor: Weston-in-Gordano, 28 September 1903, J.W.W. [det. J.G.] (BM!); thine, 4 June 1915, CUS derives: June 1917, [det. G.O.A.] (BM!); ST443731 ditch No.- 199, 6 April 1989, N.F.S., S.W., E.J.M. (SLBD; ST442733 ditch No. 195, c.1993 (S. Wilson in Jitt.); ST442733 ditch No. 195; ST443731 “ditch Nomels9: ST442733 ditch No. 195, 20 September 1999, N.F.S. et al.; ST444732 ditch No. 198, c.1990 (S. Wilson in litt.); ST444732 ditch No. 198, c.1993 (S. Wilson in itt.); ST444732 ditch No. 198; ST443731 ditch No. 199, 24 July 1995, S.W. [det. N.F.S.]; ST443731 ditch No. 199; ST445734 Bass Pond, 3 October 2001, N.F.S. et al.; 11 June 2002, N.F.S. Chara aspera Deth. ex Willd. Rough Stonewort White (1887) originally assigned several records to this species, but he subsequently corrected this, saying “I now know that the plants named C. aspera in my earlier book [refers to White 1887] should have been placed CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET 151 [in C. hispida]|” (White 1912). The only record from v.c. 6 which may represent this species is recorded as “probably this” from the [Bridg- water and Taunton] canal near Bridgwater, again it has not been possible to confirm this record and this species can not be confidently recorded for v.c. 6. V.C. 6 BEFORE 1994 [ST23 - “Probably this” canal, near Bridgwater (Marshall 1914)]. Chara contraria Kitz. Opposite Stonewort Synonyms: C. vulgaris var. contraria (A. Braun ex Kitz.) J. A. Moore, C. vulgaris var. hispidula (A. Braun) J. A. Moore C. contraria is probably under-recorded in the region; all records from v.c. 34 were made since 1994 by R.V.L. and determined by N.F.S. It is likely that before 1994 this species was overlooked for C. vulgaris. C. contraria appears to be widespread but local with populations in calcareous lakes, flooded gravel pits and ditches. It often grows in relatively shallow water at the edge of the waterbodies, typically dying back as water levels fall in summer, to reappear as a dense sward following the first rains of the autumn. Until recently, it was considered nationally scarce (Stewart & Church 1992), but has recently been shown to be more widespread than previously thought (Stewart 2004) and locally frequent in the Thames catchment and East Anglia. V.C. 6 BEFORE 1994 ST33 - (As C. vulgaris var. contraria) ST385301 North Moor, Othery, 1986, D.R. [conf. J.A.M.] (BM!). ST47 - Rhine below Clapton-in-Gordano, 1922, C.LS. [det. G.R.B.-W.] (White 1923; Willis & Jefferies 1959). 1994 ONWARDS ST45 - ST436540 rather rare, NW corner of Cheddar Reservoir, 3 October 2001, N.E.S.; ST441537, 18 September 2004, B.G. et al. [det. N.F.S.] (BM!). ST55 - ST515596 Blagdon Lake, 19 Sept- ember 2003, B.G. et al. [det. N.FS.] (BM!). - ST569597 Chew Valley Lake, 20 September 2003, B.G. et al. [det. N.F.S.] (BM!). V.C. 34 1994 ONWARDS SO70 - SO753076 flooded, disused gravel pit, Frampton on Severn, 2 September 1995, RVG! deta Nekesal: - §$0750090 flooded, disused gravel pit, Saul, 2 September 1995, R.V.L [det. N.F.S.] (RVL!). - $O752092, small pond northeast of main gravel pit, Saul, 2 September 1995, R.V.L! (det. N.F.S.]. ST78 - ST717887 deep quarry pool north of Wickwar, 1995, R.V.L! [det. N.F.S.]. ST80 - ST813015 upper (western), large artificial lake, Woodchester Park, 26 June 1995, R.V.L [det. N.F.S.] (RVL!). ST89 - ST899986 Cherington Park Lake, 1995 R.V.L! [det. N.F.S.]. Chara globularis Thuill. Fragile Stonewort Synonyms: C. globularis Thuill. var. capillacea (Thuill.) Zaneveld.; C. globularis Thuill. var. hedwigii (Agardh. ex Bruzelius) J. A. Moore. C. globularis is widespread and fairly frequent in Britain. Within the region it was recorded from a wide variety of waterbodies. Some records prior to Groves and Bullock-Webster (1924) probably refer to C. virgata which had previously been separated only at varietal level. This particularly applies to records given as var. capillacea which H. and J. Groves usually applied to specimens now included in C. virgata. Some of the apparent decline may be due to a lack of survey or a lack of identification of charophytes to species level during surveys of the Somerset Levels com- bined with changes in our understanding of the taxonomy of this and C. virgata. However, it does appear to have been genuinely lost from a number of sites and if this loss can be confirmed, measures should be _ identified which could restore populations. V.C. 6 BEFORE 1994 ST25 - ST295517 pool, Berrow sand dunes, 26 June 1980, M.V.M. [det. J.A.M.] (BM). ST33 - Bridgwater and Taunton Canal, H.S.T. (Murray 1896). ST34 - In deep water of excavated scrape, Tealham Moor, 1992, R.S.C. (Willis 1992). ST44 - Turf pits near Ashcott railway station, c. 1902, C.B., D.F., J.W.W. (Bucknall et al. 1903); pits on the peat between Ashcott and Shapwick Stations (as var. capillacea) (White 1912; Marshall 1914). ST46 - Kenn Moor (White 1912; Marshall 1914). ST47 - Near Clevedon, 1883, W.J. (Groves & Groves 1886; Murray 1896; White 1912). - Ditches, Clapton Moor, c. 1902, C.B., D.F., J.W.W. (Bucknall et al. 1903). 152 ST76 - In canal basins [Bath] (as C. hedwigii) (Murray 1896; White 1912). 1994 ONWARDS ST33 - Southlake Moor: ST36723054, northern end of ditch No. 78 , 20 July 2000, NEES: ICR 4) October 20012 NEES: ST36813052, ditch No. 79, 1 October 2002, N.F.S.; ST37043045, ditch No. 99N, 4 October 2001, N.F.S.; ST36903034, ditch No. 137E, 23 October 2001, N.F.S. ST44 - ST414426 ditch north-west of Westhay Level Peat Works, July 1995, S.G! ST45 - ST441537 Cheddar Reservoir, 18 September 2004, B.G. et al. (BM!). ST47 - ST442736, 1997, R.V.L. (RVL!). - Gordano Valley: - Walton Moor: 1902 (as_ var. capillacea) (White 1912; Marshall 1914; Willis & Jefferies 1959); ST432727 ditch (No. 46), 5 m altitude, 4 March 1989, N.E.S., E.J.M. (SLBD; ST425725 ditch No. 8; ST426725 ditch No. 11; ST430726 ditch No. 20; ST430726 ditch No. 29, 6 April 1989, N.E.S., S.W., E.J.M.; ST431727 ditch No. 46, 4 March 1989, N.F.S., E.J.M. (Willis 1989); ST428724 ditch No. 19, 20 September 1999, N.F.S. et al.; ST433730 S end of ditch No. 97, 20 October 2002, N.E.S et al.; ST434733 ditch No. 101, 11 June 2002, N.EF.S. - Weston Moor: ST442735 ditch No. 179; ST443736 ditch No. 180; ST443735 ditch No. 186, 6 April 1989, N.F.S., S.W., E.J.M. [conf. J.A.M. 1989] (SLBDI) (Willis 1989); ST442735 ditch No. 179, 20 September 1999, N.F.S. et al.; ST442737 ditch No. 148, 3 October 2001, N.F.S. et al.; ST435731 ditch, 11 June 2002, N.ES.; ST441737 ditch No. 160-162, 24 February 2004, N.E.S. S1Sd4- Sis l5596 ‘Blasdons ake, 919 September 2003, B. Goldsmith er al. [det. N.EF.S.] (BM!). - ST569597 Chew Valley Lake, 20 September 2003, B. Goldsmith et al. [det. N.E.S.] (BM!). V.C. 34 BEFORE 1994 SO60 - SO620089 neglected lake, dredged c. 10 years ago, Brock Lane, Parkend, 20 June 1984, M.V.M. [det. J.A.M.] (Bailey & Moore 1985; Moore & Bailey 1986). SO61 - S0O662110 pond, Soudley Brook, Forest of Dean, June 1975, D.N.A., M.J.A., S.C.H. [det. J.A.M.] (Moore & Bailey. 1986). R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN SO90 - Canal, Chalford (Riddelsdell et al. 1948). - Pond, Rudgeway, Alveston, 1925 C.T. (White 1926). ST68 - Froglane Pit, Coalpit Heath, 1955, G.W.G. [det. G.O.A.] (Sandwith & Sandwith 1956; Moore & Bailey 1986). ST78 - Wickwar — Yate Area: - [ST776848] field pond near southwest corner of Bodkin Hazel Wood, nr. Horton, 1952, G.W.G. [det. G.O.A.] (Sandwith & Sandwith 1953; Moore & Bailey 1986 as var. capillacea). - Yate Lower Common in a strontium pit, H.J.R. (Riddelsdell et al. 1948). - Wickwar area, 1920, J.W.W. (BM); 1917, J.W.W. (LSR) (White 1919; Riddelsdell et al. 1948; Moore & Bailey 1986). - $T108857 strontia pit between Yate Court and Leech Pool Farm, Yate, 15 December 1984, A.J.B. [det. J.A.M.] (Bailey & Moore 1985; Moore & Bailey 1986). - Barbours Court Farm, Wickwar, 10 December 1990, N.F.S. , M.A.R.K., C.K. ST89 - near White Horse, Avening, 3 July 1926, H.J.T. (BM!). 1994 ONWARDS SO70 - > SO753076 flooded Weravely pie Frampton-on-Severn, 2 September 1995, R.V.L [det. N.F.S.] (RVL!). - $0748076 pond, Frampton on Severn, 1977, A.K. [det. J.A.M.] (Moore & Bailey 1986); SO753076 flooded gravel pit, 2 September 1995, R.V.L. (RVL!); SO754077 flooded gravel pit, 1995 R.V.L. (RVL!). - S§0733056 ditch, New Grounds, Slimbridge, 29 August 1977, A.K. [det. J.A.M.] (Moore & Bailey 1986); SO720048, 1996 R.V.L. Chara hispida L. Bristly stonewort C. hispida is sparsely distributed in Britain, becoming rarer northwards and westwards. It has been recorded from a variety of calcareous habitats in the region, most frequently in the Gordano Valley and Somerset Levels, with other scattered records from ponds and larger waterbodies. In 1996, an _ unidentified stonewort, probably this species, was recorded from ponds in the National Trust Property at Westbury-on-Severn, but subsequent visits in 1997, 2000 and 2002 have failed to find any stoneworts. CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET 153 This species appears to have declined throughout the region. In v.c. 6, it has only been recorded since 1994 from the Gordano Valley although before 1994 it was apparently widespread in the Somerset Levels. In v.c. 34, it has never been abundant and has only been recorded from scattered pools in the Forest of Dean. Few of these have been surveyed for charophytes in recent years and it is possible that a reasonably healthy population remains in the area. V.C. 6 BEFORE 1994 ST33 - Ditch, near Weston Zoyland, Kings Sedge Moor, 23 August 1907, E.S.M. (BM!, CGE!) (Marshall 1914; 1908). - Nr. Middlezoy, W.W. (Marshall 1914). - Nr. Othery, 1909; 1914). - Drain near Sedgemoor Cut, May 1899, G.R.B.-W. (Bullock-Webster 1901). ST45 - Ditches, Clevedon (White 1887 as C. aspera, Murray 1896; White 1912, Marshall 1914). ST46 - Marsh ditch Kenn Moor, 28 September 1903, J.W.W. (CGE!); peaty ditch Kenn Moor 17 September 1904, J.W.W. [det. A.B.] (BEL!, BM!, CGE [det. J.G.]!, E!, NMW!, OXF!, SLBI!), 17 September 1904, J.W.W. [det. A.B.] (CGE [det. J.G.]!, NM'W!) (Groves 1905; White 1912, Marshall 1914). - Ditches Yatton (White 1887 as C. aspera; Murray 1896; White 1912; Marshall 1914). ST47 - Ditches, Portbury (White 1887 as C aspera; Murray 1896; White 1912; Marshall 1914 gives recorder as E.G.). - Nailsea Moor, 4 September 1902, J.W.W. (CGE!). - Gordano Valley: - peaty ditch Clapton Moor, 28 September 1903, J.W.W. (CGE!, NMW!, OXF!) (White 1920); ditch, peaty meadow, 28 September 1903, J.W.W. [conf. G.R.B.- W.) (E!) (White 1912). Siiseesesi7 05322 Park Wood, south ‘of Redlynch, August 1984, J.G.K _ [det. J.A.M.] (BRC). ST76 - “In the canal” [Bath] (Babington 1834; White 1887; 1912); canal between Bath and Bathampton, 4 July 1956, J.P.M.B. (BM). G.R.B.-W. (Marshall 1994 ONWARDS ST47_ - Gordano Valley: - Walton Moor: 1867, J.D.H. (BM) (White 1887; Murray 1896; White 1912; Marshall 1914); 25 June 1849, C.C.B. (CGE!); 13 August 1922 F.G. (LSR) (Willis & Jefferies 1959); ST444735 ditches, 7 May 1984, A.L. [det. J.A.M.] (BM!); ST430730 ditch (No. 137), 5 m altitude, 4 March 1989, N.F.S., E.J.M. (SLBI); ST426725 ditch No. 11; ST428724 ditch No. 19; ST430726 ditch No. 20; ST429725 ditch No. 22; ST430726 ditch No. 29, 6 April 1989, N.F.S., S.W., E.J.M.; ST427725 ditch No. 18/18.1; ST429725 ditch No. 22; ST430726 ditch No. 29, 20 September 1999, N.F.S. et al.; ST434733 ditch? Nov 10 ft June” 20027 NES: ST434727 ditch No. 53, 20 October 2002, N.ES et al. - Weston Moor: 1903 G.C.D. (OXF!); ST443735 ditch No. 186; ST443731 ditch Now 199350) April” 1989e aN Ss SAW... E.J.M.; ST443736 ditch No. 180, 30 June 1989, S.W., O.M., [det. J.A.B.]; ST442735, 1997, R.V.L. (RVL!); ST442734 ditch No. 195556 —June> 1999" ONE'S” RV: ST442735 ditch No. 179; ST442733 ditch Nos 19537 Si443731 ditch No: 199)" 20 September 1999, N.F.S. et al.; ST442739 ditch No. 169 [det N.F.S.]; ST443731- ST444732, ditch No. 199, 3 November 1999 A.W!; ST440738 ditch No. 159, 24 February 2004, N.F.S.; ST438736 ditch No. 154/155; ST442738 ditch No. 166-168, 11 June 2002, N.F.S.; ST441734 ditch No. 176; ST441735 ditch No. 176A; ST442735 ditch No. 179, 6 June 1999, N.F.S., R.V.L.; ST441734 ditch No. 184; ST443731 ditch NOS 199 Si 444732 ditch No: > 206: ST445734 Bass Pond, 3 October 2001, N.E.S. et al.; ST445734 Bass Pond, 11 June 2002, N.F.S.; ST442734 ditch No. 193, 20 October 2002, N.E.S et al. V.C. 34 BEFORE 1994 SO61 -S0O644124 pond near Lightmoor Pond, Ruspidge, 31 August 1977 A.K. [det. J.A.M.] (BRC); Lightmoor Colliery, Ruspidge, 1980, A.O. (Moore & Bailey 1986); SP642122 Lightmoor Pond, October 1980 S.C.H. [det. J.A.M.] (BM); 1994 C.W., D.H. (English Nature 1994). SO71 - [probably this] Water garden ponds, Westbury Court, Westbury-on-Severn, 154 R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN 19735 S:@3H Kee. det RerEBalGlolland 1974, Moore & Bailey 1986). 1994 ONWARDS SO61 - Crabtree Colliery, Cinderford, 2 October 1980 A.O. (Bailey & Moore 1985, Moore & Bailey 1986); SP639134, December 1997, R.V.L! Chara virgata Kitz. Delicate stonewort Synonym: C. delicatula Agardh; C. globularis var. virgata (Kiitz.) R.D. Wood C. virgata is widespread and _ frequent throughout Britain, particularly in the west. In the region, it appears to be mainly an early colonizer of new waterbodies, particularly worked strontium pits and farm pools in the Wickwar area and it is possible that it tends to decline after a few years as a result of competition from other charophytes and vascular plants. It is still frequent in ditches in the Gordano Valley where it prefers the more peaty substrates. Whilst many of the pools from which it has been reported have been lost to development or in filled for agriculture in recent years, this is still one of the most frequently recorded charophytes in the region and there is no real evidence of a decline. Forms with well developed stipulodes and more prominent spine cells have been recorded from Shapwick and the Gordano Valley as C. delicatula Desv. var. barbata (Ganterer) J. Groves and Bull.-Webst. V.C. 6 BEFORE 1994 ST43 - Moor ditch below Compton Dundon, 14 July 1891, R.P.M. (as C. fragilis) (BM!, MANCH!) (Murray 1896). ST44 - Shapwick Moor, 10 May 1899, G.R.B.-W. (CGE); Shapwick, 1953, A.J.D. (as C. delicatula var. barbata), [det. G.O.A.] (Sandwith & Sandwith 1954, Wallace 1954). - Pond near Shapwick Station, 8 August 1937, J.P.M.B., C.I.S. (BM). ST47 - Nailsea Moor (White 1912 as C. globularis, Marshall 1914); 10 September 192 I (as; Garages) eS le CAS ez val: (BM!). 1994 ONWARDS ST47 - Gordano Valley: rhines, 1925, [det. J.G.] (White 1926, Willis & Jefferies 1959) - Walton Moor: ST431731 ditch No. 138, Walton Moor, 5 m altitude, 4 March 1989, N.F.S., E.J.M. (SLBI) (Willis 1989); ST425725 ditch No. 8; ST426725 ditch No: ~ 11 (Willis 1989); ST428724-ST427724, ditch No. 19 (Willis 1989); ST427724— S427725 ditch No. 18/18-1 (Willis 1989); ST429725 ditch No. 22, 6 April 1989, N.F.S., S.W., E.J.M.; ST427725 ditch No. 18/18.1 [det. J.A.B.]; ST428724 ditch No. 19, 29 June 1989, S.W., O.M., [det. J.A.B.]; ST436734 ditch near 107, 30 June 1989, S.W., O.M., [det. J.A.B.]; ST425726 ditch No. 6, 11 March 1992, S.W. [det. J.A.B.] (in J. A. Bryant correspondence file); ST427725 ditch No. 18/18.1, 20 September 1999, N.E.S. et al.; ST434727 ditch No. 53; ST435727 ditch No. 54, 20 October 2002, N.ES. et al. - Weston Moor: Weston-in-Gordano (as C. delicatula var. barbata), 1903, G.C.D. (OXF!) (Groves & Bullock-Webster 1924); ST444735, 7 May 1984, A.L. [det. J.A.M.] (BM); ST438734 peaty ditch No. 171, Weston Moor, 5 m altitude, 5 March 1989, N.F.S. (SLBI) (Willis 1989); ST443737 ditch No., 147 (Willis 1989); ST442738 ditch No. 167 (Willis 1989); ST443737- ST443738, ditch No. 168 (Willis 1989); ST439734 ditch No. 171N; ST442735 ditch No. 177 (Willis 1989); ST442735, ditch No. 179, 6 April 1989, N.F:S., S.-W; EM. (Willis 1989); ST443736 ditch No. 180, 30 June 1989, S.W., O.M.; [det. JACB:I: ST442734, ditch No. 193, 11 March 1992, S.W. [det. J.A.B.] (Gn JP VAgieBryant correspondence file); ST442736 Weston Moor, 15 May 1997 R.V.L. (RVL!); ST441734 ditch No. 176; ST441736 ditch No. 177; .ST442735 ditch” Nowa: ST443731 ditch No. 199, 6 June 1999, N.E‘S., R.V.L.; S1T442734) ditchsNow iss: ST442733 ditch No. 195; ST444732 ditch No. 206, 20 September 1999, N.F‘S. et al.; ST441734 ditch No. 184; ST442734 ditch No. 193; S1444732 ditch )Noww206: ST445734 Bass Pond, 3 October 2001, N.ES. et al.; ST441737 ditch No. 160-162; ST446733 W of ditch No. 212.1; ST445734 Bass Pond; ST445733 Pool S of Bass Pond, 11 June 2002, N.F.S.; ST442734 ditch No. 193, 20 October 2002; NES tera: ST438738 ditch No. 121-124; ST441737 ditch No. 160-162, 24 February 2004, N.E:S. - Ditch, Clapton Moor ST454735, 3 November 1999, A.W!; rare, ditch, Clapton Moor ST460734, 3 October 2001, N.F.S. et al. - §T52503611 Butt Moor, ditch No. 1, 23 July 2003, N.F.S, E.J.M. et al. CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET (I5)5) V.C. 34 BEFORE 1994 SO68 - Wickwar — Yate area: — Pond, Coal Pit Heaths > Eroglane; 21 September 19110; J.W.W. (CGE) (Riddelsdell et al. 1948 — as Coalpit Heath is probably this record). SO6878 - Kingsgate Park, by Stanshawe’s Court, Yate, 1978, G.W.G. (Willis 1978, Moore & Bailey 1986). SO80 - SO8902 Thames and Severn Canal (Riddelsdell et al. 1948). SO90 - SO9002 Thames and Severn Canal (Riddelsdell et al. 1948). ST78 - Wickwar — Yate area: — various times in at least seven ponds, including Goosegreen Farm, nr. Yate [det. H. & J.G.] (White 1912; Riddelsdell et al. 1948). - pond, north east of Lance Coppice, Inglestone Common, 1953, G.W.G. (Moore & Bailey 1986) - strontia pits, Goose Green, Yate, 1953, G.W.G. (Moore & Bailey 1986) - standing water on old mineral railway, Froglane, Mayshill, 1954, G.W.G. (Moore & Bailey 1986) = mill pond, Whitwell Bottom, Lower Kilcot, 1955, G.W.G. (Moore & Bailey 1986) - pond on the skirt of Yate Lower Common, south of Hall End, 4 June 1917, J.W.W. (as C. fragilis) (RNG!). Seevaici O07 18825, 7 May H.J.M.B! (RNG!). - pool in an old clay pit between Rangeworthy and Hall End, 16 July 1910 J.W.W. [det. H. & J.G.] (Moss 1911). ST96_ - Gall Pond, Tortworth, A.J.B. 1955 1994 ONWARDS SO70 - SO750091 flooded gravel pit, Saul, 2 September 1995, R.V.L! SO80 - SO816015 old pond, Woodchester Park, 28 August 1995 R.V.L. ST78 - Wickwar — Yate area: - Yate Court, 1954, (Moore & Bailey 1986); ST710860 Yate Court, 1995, R.V.L! - $T702884 [Newland’s Farm], 14 June 1978 R.H.B., J.A.M. (BM) (Willis 1978, Moore & Bailey 1986); ST703883, 10 February 1990 N.F.S., M.A.R.K., C.K.; ST702883, 19 August 1995 R.V.L!; ST702884, 19 August 1995, R.V.L!; ST703883, 19 August 1995 R.V.L. - ST710859, 19 August 1995, R.V.L! ST89 -ST870992 Lake, Longford’s Mill, near Nailsworth, 9 June 1997, R.V.L. Chara vulgaris L. Common Stonewort C. vulgaris is frequent in Britain, particularly in the lowlands. It is the most widespread and frequently recorded stonewort in the region. It occurs in most types of waterbody, even in shallow, still backwaters of flowing rivers. It appears very capable of colonising newly created ponds, particularly in lime-rich areas, tolerating some competition from _ other charophytes and vascular plants such as Lemna minor (Common Duckweed), Zannichellia palustris (Horned Pondweed) and Elodea canadensis (Canadian Waterweed). Within the region, it probably has the most stable population size of all the stoneworts, colonising new waterbodies as other populations decline and disappear. Although there are fewer records in the region since 1994 than before, this is likely to be mainly due to a lack of survey and, when the abundance of the three common varieties is taken into consideration, there is no evidence to suggest that it is declining. No detail is given of the status of the varieties of C. vulgaris; the varieties tend only to be identified occasionally and almost never on a systematic basis. It is therefore difficult to interpret apparent changes. It is likely that many of the records listed as C. vulgaris s.1. refer to C. vulgaris var. vulgaris however this was not specified on information relating to the record. V.C. 6 BEFORE 1994 ST23 - Bridgwater, H.S.T. (Murray 1896). ST2535 - Pond, Brean, 3 August 1882, R.P.M. (BM) (Murray 1896); 16 September 1906, H.S.T. (BM). ST33 - Weston Zoyland, (Marshall 1908; 1914). - §T370305 Burrow Mount, 12 July 1987, M.A.R.K., C.K. ST34 - Near Edington Burtle, 8 June 1835, (possibly also var. longibracteata referred to as recorded by W.B.W. in White 1912) T.C. (BM) (White 1912; Marshall 1914). - Ditch, Edington Junction, 23 August 1915, E.S.M. (BM). - Mark Moor, 1915, C.I.S. [det. G.O.A.] (Sandwith & Sandwith 1958). - Burnham, W.B.W. (White 1912). ST36 - ST3868 Rhine, Kingston Seymour, 16 April 1981, P.M.W. (PMW!). - ST345610 ditch, north of Weston Airport, Weston-Super-Mare, 1990, J.N! 1907, E.S.M. 156 R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN - $T383638 rhyne near West Hewish, 17 February 1990, M.A.R.K., C.K! - §T392604 southern side of track near Woolvershill Batch, Banwell, 17 February 1990, M.A.R.K., C.K! ST43 - King’s Sedge Moor, 13 August 1883, R.P.M. (BM). ST44 - ST4145 Tadham Moor, 4 June 1979, M.A.P. (BRC). - Ditch near Catcott Burtle, 3 August 1937, J.P.M.B., C.I.S. (BM). ST45 - Clevedon, D.F. (Murray 1896; White 1912). ST46 - Moor ditches, Yatton (White 1912). - Kenn Moor, 11 October 1953, C.LS., N.Y.S. [det. G.O.A.] (BM) (Sandwith 1962). ST47 - Nailsea Moor, 1949, C.LS. [det. G.O.A.] (Wallace 1950); ditch on Nailsea Moor, 27 July 1914, J.W.W. [det. J.G.] (Britton 1918). ST53 - Kingweston (Murray 1896). ST56 - Reservoir, Barrow Gurney, 6 October 1933, H.S.T. (BM). - Quarry pool, Hartcliff Rocks, I.M.R. (White 1912; Marshall 1914). ST63 - Pond, Bruton, September 1936, F.K.M. (BM). ST65 - Old coal canal, Camerton (White 1912). ST73 - Roadside pond near Penselwood, 20 July 1891 R.P.M. (BM) (Murray 1896). ST76 - “In the canal” Bath (Babington 1834; Murray 1896). 1994 ONWARDS ST25 - Dune slacks near Berrow (White 1912; Marshall 1914); [ST2952] pond, 20 May 1929, J.E.L. (BM); 20 September 1950, C.I.S. (BM); Pool west of church, 1992 R.S.C. (Willis 1992). Si53) <2 Ditches aisiouthliakes Moon: ST36393048, ditch No. 53; ST374302 ditch No. 1208 June i998 Ravel Sale KEP LiWe: S136243055." editch FaNow 1: S363 1380572 ditch Nom 32 Si36733050; ditch No. 58; ST36913057, ditch No. 61; S1367230545) ditch Now) 734 NN iend: ST37003053, ditch No. 81; ST36813037, ditch No. 95; ST36853038, ditch No. 96; ST36983045, ditch No. 98; ST37043045, ditch No. 99N, 20 July 2000, N.F.S., K.P.; ST36193025, ditch No. 64; ST36333026, ditch No. 85, 26 July 2000, N.F.S., P.W., K-P.; S136723054, ditch No. 78, N end; ST36983045, ditch No. 98; ST37043045, ditch No. 99N, 4 October 2001, N.F‘S.; ST36373030, ditch No. 88; ST36903034, ditch No. 137E; ST36783030, ditch No. 147 23;) eOctober ) 20015. 3NeEsSe ST36313027, ditch No. 86; ST37413027, ditch No. 120; ST36903034, ditch No. 137E; ST37062984, ditch No. 194 19, September 2003, N.F.S. - ST308392 roadside ditch in N. edge of Bridgwater, 26 July 2004, N.FS. ST42 - ST483260 trial pit, in Appledoor Quarry, 10 May 2002, M.R.H! ST44 - Tealham Moor: ST40794520 ditch No. 1, 9 May 2003, N.F.S.; ST40644540 ditch No. 3, 23 February 2004, N.ES:.; ST40504521 ditch No. 6, 23 February 2004, N.F.S.; ST40804520 ditch No. 10, 9 May 2003, N.F.S.; ST40794509 ditch No. 11, 10 June 2004, N.F.S.; ST40644524 ditch No. 12, 9 May —200355NiIE Ss. ST40924505 ditch No. 13, 9 May 2003, N.F.S.; ST41024515 ditch No. 14, 9 May 2003, N.F.S.; ST41084527 ditch No. 15, 9 May 2003, N.F.S.; ST41144520 ditch No. 16, 9 May 2003, N.F.S. ST47 - Gordano Valley: 1914, C.I.S. [det. G.0.A.]. - Walton Moor: ST430730 ditch No. 137, 4 March 1989, N.F.S., E.J.M. - Weston Moor: ST439734 ditch No. 171N; ST443736 ditch No. 180, 5 March 1989, N.F.S.; ST441736 ditch No. 177; ST442735 ditch No. 179; ST442737 ditch No. 148; ST442738 ditch No. 166-168; ST443735 ditch No. 186; ST443736 ditch No. 180;.ST443737 ditch No. 147, 6 April 1989, N.E.S., S.W., E.J.M.; ST439734 ditch No. 171N; ST443736 ditch No. 180, 30 June 1989, S.W., O.M. [det J.A.B]; ST443731 ditch No. 199, 6 June 1999, N.ES., R.V.L.; S1T445737 ‘ditchiNomiSirys November 1999, [det N.F.S.]; ST443731 ditch No. 199, 20 September 1999, N.F.S. et al.; ST438736 ditch No. 154/155; ST438738 ditch No. 121-124; ST442738 ditch No. 166-168; ST443737 by E end of ditch No. 147; ST444736 ditch No. 192; ST444738 ditch No. 181; ST445734 Bass Pond, 11 June 2002; ST446733 ditch No. W of 212.1, 11 June ~2Z0025s Naas ST438738 ditch No. 121-124, 24 February 2004, N.F.S. - $T493738 Cattle trough near Downs School, 2002, P.M! ST53 - Butt Moor: ST52563606 ditch No. 2, 26 July 2000, N.F.S., P.W.; ST52563606 CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET sq) ditch No. 2, 23 October 2001, N.F.S.; ST52623609 ditch No. 3; ST52613600 ditch No. 4; ST52573614 ditch No. 5; ST52673614 ditch No. 6, 10 October 2002, NEERSes eS ls2503611" “ditch No. — 1; ST52503585 ditch No. 8; ST52563606 ditch No. 2; ST52223608 ditch No. 11; ST52173610 ditch No. 12; ST52213615 ditch No. 13, 23 July 2003, N.F.S., E.J.M. Camas SlS25138589" ditch” No.7; ST52463583 ditch No. 9; ST52663563 ditch No. 10, 17 September 2003, N.F.S. ST55 - ST515596 Blagdon Lake, 19 Sept- ember 2003, B. Goldsmith et al. [det. N.F.S.] (BM!). ST58 - ST530807 New mitigation ponds, Avonmouth, 1999, V.H! V.C. 34 BEFORE 1994 SO50 - SO565065 Clearwell Quarry, 27 July 1986, S.H.B, S.C.H. [det. J.A.M.] (BRC). SO51 - Jugshole Pool, Coleford, 1980, S.C.H., M.J.A. [det. A.O.] (Moore & Bailey 1986). SO60 - SO612033 lake, Lydney Park, 1980, A.K. [det. J.A.M.] (Moore & Bailey 1986). _ = pond in marshy field by River Severn, Poulton Court Farm, Awre, 1988, S.C.H. [det. J.A.M.]. SO61 - Speech House (Riddelsd. et al. 1948). - $0695067 Shallow pools on track, Lennetshill Plantation, Cinderford, 1980, A.O. (Bailey & Moore 1985). - Shallow calcareous pool and ditch, Crabtree Plantation, Cinderford, 1980, A.O. (Bailey & Moore 1985). - Soudley valley, H. (Riddelsdell ez al. 1948); SP662116 Upper Soudley pool, August 1976, S.C.H. [det. J.A.M.] (BRC); 1994 C.W., D.H. (English Nature 1994); 1994, C.W., D.H. (English Nature 1994). SO70 - SO733056 ditch, New Grounds, Slimbridge, 29 August 1977, A.K. [det. J.A.M.] (BRC). - Nupend (Riddelsdell et al. 1948). SO71 - Highnam (Riddelsdell et al. 1948). - Arlingham (Riddelsdell et al. 1948). SO72 - Pond near Newent Canal (as C. hispida), G.A.O.B. (Riddelsdell et al. 1948). SO80 - Woodchester Park, H.P.R. (Riddelsdell et al. 1948). SO90 - Pool by canal above Chalford, H.P.R. (Riddelsdell et al. 1948). ST57 - Quarry pools, Eastfield, Westbury-on- Trym (White 1887; 1912). - Ponds, Durdham Down Shirehampton (White 1912). and ST67 - Brickworks ponds, Shortwood, Mangotsfield, 1955, G.W.G. (Moore & Bailey 1986); 28 October 1964, D.M.-S. [det. S.P.P.] (Moore & Bailey 1985). - Duchess Ponds, Stapleton (White 1912). - Ditch, Siston Common, 1882, D.Ha. (White 1887; 1912); 1920 D.Ha. (White 1912). ST68 - Standing water on old railway, Froglane, Mayshill, 1964, G.W.G. (Moore & Bailey 1986). - Woodlands, Patchway (Riddelsdell et al. 1948). ST69 - ST633977 newly dug pond north of Hill Pill, 16 November 1986, M.A.R.K., C.K!; ST628973 newly dug pond, Hill Pill, 16 November 1986, M.A.R.K., C.K! ST75 - [ST7159] canal near Dunkerton (as C. contraria), 25 June 1910, C.B., J.W.W. [det. H. & J.G.] (BM, MANCH! specimen labelled C. fragilis probably misprint) (Moss 1911; White 1912; Marshall 1914). ST78 - Wickwar — Yate area: ST791888 mill pond, Whitewell Bottom, Lower Kilcot (Riddelsdell et al. 1948); ST7183, 1910, J.W.W. (CGE! - probably the same record, located by C.B. and preserved by J.W.W.); 1953, (Moore & Bailey 1986); 1955, G.W.G. (Moore & Bailey 1986). ST79 - ST786932 sunny pond, Newark Park Estate, May 1984, M.S., W.L. (Clements et al. 1985); ST786932 pond, Newark Park, 16 May 1985, W.L. [det. J.A.M.] (BRC). ST88 - Ornamental Lake, Westonbirt School, 1984, R.H.B. (Bailey & Moore 1985, Moore & Bailey 1986). ST89 - Cherington Park Lake (Riddelsdell et al. 1948); ST898986, 7 July 1977, R.H.B., J.A.M. [det.- J.A.M.] (BM) (Moore & Bailey 1986). - Pond, Midland Fishery, Nailsworth, 2 September 1949, D.F.L. (BM) (Moore & Bailey 1986). - Pond near White Horse, Avening, 3 July 1926, H.J.R. (as C. fragilis) (BM!) (Riddelsdell et al. 1948). I.M.R. 1994 ONWARDS SO61 - S0O639134 Crabtree Colliery, Cinderford, December 1997, R.V.L! SO70 - Frampton Pools SSSI, 1993, C.W., A.H. (English Nature 1994); Frampton Sailing Lake, 1993, C.W., A.H. (English Nature 1994); SO753076 flooded gravel pit, 1995, R.V.L. - $0750090 flooded gravel pit, Saul, 1995, R.V.L. 158 R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN SO72 - SO713266 pond nr. August 1998, R.V.L. SO80 - Thames and Severn Canal: SO858042 Brimscombe, 27 July 1997, R.V.L.; SO848051 Stroud, 1997, R.V.L.; SO8104 Ebley, 7 October 2003 R.V.L. - §$085810203 small, spring-fed pool used by cattle, Black Ditch, edge of Minchinhampton Common, 14 January 2004, D.C. SO90 - SO967004 abundant in Severn and Thames Canal near Coates, 1997, R.V.L. (RVL!) ST69 - ST674976 pond, Whitcliff Deer Park, near Ham, 1997, M.A.R.K., C.K., B.A.G. ST78 - Wickwar — Yate area: - $T710859 Yate Court, 1954, (Moore & Bailey 1986); - ST716888 margin of deep quarry pool, 995, ROVE - ST703883 pond, Newland’s Farm, OOS Reale - ST757883 pond east of Lance Coppice, 1953, G.W.G. (Moore & Bailey 1986); 1995, R.V.L.; 1996, R2V-L.; 1997, R:V.E.; 1998; REV.E.; > June’ 19995 N-E:S., R.V.L. ST79_ - ST771939 lake, Tyley Bottom, 1978, R.H.B., J.A.M. [det. J.A.M.] (Moore & Bailey 1986); ornamental lake, Tyley Bottom, 1984, R.H.B., J.A.M. (Bailey & Moore 1985); ST775942 recently dug farm pond, Tyley Bottom, 1996, R.V.L. ST89_ - ST803973 new pond at the top of the stream entering Owlpen Valley, 1997, R.V.L.; newly created pond on site of old dew pond on farm adjacent to Owlpen Manor, 2003, R.W. (supplied by M. Palfrey) [det. M.A.R.K]. - Marl pool, Ruskin Mill, 1 June 1996, J.J.D! Newent, 31 var. crassicaulis (Schleich. ex A. Braun) Kitz. V.C. 6 or 34 (the vice county boundary runs through the middle of the reservoir) BEFORE 1994 ST77 - Monks Wood Reservoir, Bath, 1928 C.LS. [det. J.G.] (White 1928). var. longibracteata (Kitz.) J. Groves and Bull.- Webst. V.C. 6 BEFORE 1994 ST47 - Gordano Valley: - Walton Moor: ST430730 ditch No. 137, 5 m altitude, 4 March 1989, N.ES.,- E.J.M. (SLBD. - Weston Moor: ST445737 3 November 1999, A.W!; ST442738 ditch No. 166-168, 6. ‘April 1989) | NUESe) Si Wee Baie ST442735 ditch No. 179, 6 April 1989, N.E.S., S.W., E.J.M. ST57 - Pond between Bedminster and Whitchurch, 1882 (White 1887; Murray 1896; White 1912). 1994 onwards ST33 - Southlake Moor: ST36933040, ditch No. 97 20 July 2000, N.F.S., K.P. ST44 - Tealham Moor: ST40604551 ditch No. 8; ST40704545 ditch No. 9, 23 February 2004, N.F.S. ; ST40794509 ditch No. 11, 9 May 2003, N.EF.S. - ditch Clapham Moor ST454735, 3 November 1999 A.W! V.C. 34 BEFORE 1994 SO70_ - Frampton on Severn (Riddelsdell et al. 1948); SO748071 at edge of Blue Pool, Frampton Pools, 25 June 1976, J.A.M. (Moore & Bailey 1978); SO748076 pond, 29 August 1977, A.K. [det. J.A.M.] (Moore & Bailey 1986). ST66 - Pond, Bitton, 18 June 1914, LMLR. [det. J.G.] (BM) (Riddelsdell et al. 1948). ST67 - Wickwar — Yate area: Newly dug ditch, Siston Common, 2 October 1962, D.M.-S. [det. S.P.P.] (Bailey & Moore 1985, Moore & Bailey 1986). ST78 - Wickwar — Yate area: - Ponds, Little Sodbury End, Sodbury Common, 1953, G.W.G. (Sandwith & Sandwith 1954, Moore & Bailey 1986). - Inglestone Common, 1953, G.W.G. (Sandwith & Sandwith 1954). - Horton, 1953, G.W.G. (Sandwith & Sandwith 1954, Moore & Bailey 1986). 1994 ONWARDS SO80 - SO816015 ornamental lake, Wood- chester Park, 28 August 1995, R.V.L. ST69 -ST638931 rhyne near Duckhole, 1996, R.V.L. (RVL). ST78 - Wickwar — Yate area: - §1T743851, one stem, pond by Vinney’s Lane, 5 June 1999, N.F.S., R.V.L. - §T729835 newly dug ornamental ponds, Yate Golf Course, 1997, R.V.L. ST89 - ST899986 Cherington Lake, 23 August 1995, R.V.L! var. papillata Wallr. ex A. Braun. V.C. 6 BEFORE 1994 SO60 - S0695067 [pond in marshy field by R. Severn, Poulton Court Farm, Awre] 4 June 1998, S.C.H. [de. J.A.M.] (J. A. Bryant correspondence file). CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET 159 ST23 - Clay pit, near Bridgwater, August 1899, G.R.B.-W. (Bullock-Webster 1901, Marshall 1914). ST44 - Shapwick Moor (White 1912; Mar- shall 1914). ST47_ - Nailsea Moor, 10 September and 17 September 1904, J.W.W. [det. H. & J.G] (BM, NMW ‘recorded as_ var. longibracteata!, K det. A.B!) (Roper 1918); ditch, 1914, J.W.W. [det. J.G.] (BM) (Britton 1918); 1948, C.LS. [det. G.O.A.] (Wallace 1950; Sandwith & Sandwith 1950). - Gordano Valley: - §T458736 abundant along ditch, Clapton Moor, 3 October 2001, N.F.S. et al. - Ditch between Portbury and Portishead, 1889, (White 1912; Marshall 1914). - Tickenham Moor (White 1912; Mar- shall 1914). ST55 - Blagdon Lake, 1937, H.S.T. (BM). ST56 - Ponds under Dundry Hill, 1886, [probably this specimen collected by] J.W.W. [det. H. & J.G.] (Groves & Groves 1887; White 1912). ST75 - ponds in Prior Park, nr. Bath (White 1912; Marshall 1914). 1994 onwards ST33 - ST36863056, ditch No. 60E 26 June NGOS RWI... S.J.L., K.P., L.W. - $1T394327 ditch beside drove near Sowy River, 2003, E.J.M. et al. ST47 - ST457737 Clapton Moor, 2003, P.M! - Gordano Valley: - Walton Moor, 1901, J.W.W. [det. H. & J.G.] (CGE) (White 1912; Marshall 1914); ditch, September 1935, G.O.A. (BM) (Sandwith & Sandwith 1958, Willis & Jefferies 1959); ST443736, 1989, S.W. [det. J.A.M.]; ST439734, 1989, S.W. [det. J.A.M.]; ST434733 ditch No. 101, 11 June 2002, N.E.S. - Weston Moor: ST439735 in a recently cleared peaty ditch No. 171 with some iron ochre, Weston Moor, near Weston in Gordano, 5 m altitude, 5 March 1989, N.F.S., S.W. [conf. J.A.M. 1989] (Willis 1989); ST439734 ditch No. 171N, 6 April 1989 , N.F.S., S.W., E.J.M.; ST442733 ditch No. 195, 20 September 1999, N.E.S. et al. S155 - S1569597 Chew Valley Lake, 20 September 2003, B. Goldsmith et al. [det. N.F.S.] (BM!). V.C. 34 BEFORE 1994 SO61 - Pond, Cannop, 12 July 1976, [det. J.A.M.] (BRC). SO70 - SO748071 at edge of Blue Pool, Frampton Pools, 25 June 1976, J.A.M., R.H.B. (Moore & Bailey 1977, Moore & Bailey 1978 as recorded by J.A.M.). SO80 - Thames and Severn Canal, Brims- combe, 1953, J.E.L. [det. G.O.A.] (Moore & Bailey 1986). SO90 - Lake, Misarden Park [as Miserden, however this spelling refers to the village], 1977, A.K. (Moore & Bailey 1986). - Pool by canal above Chalford, H.P.R. (Riddelsdell et al. 1948). ST58 - Water tank, Filton aerodrome, 7 October 1948, D.McC. [det. G.O.A.] (RNG!). ST78 - Wickwar — Yate area: - Ponds, Little Sodbury End, Sodbury Common, 1953, G.W.G. (Sandwith & Sandwith 1954, Moore & Bailey 1986). - Kingrove Common, Chipping Sod- bury, 1954, G.W.G. (Moore & Bailey 1986). - Inglestone Common, 1953, G.W.G. (Sandwith & Sandwith 1954) - Goosegreen Farm, Yate, C.B. (White 1912; Riddelsdell et al. 1948). - ST716817 pool on strontium-rich marl at) ic. 200) it, Yate; 1) November 1954; H.J.M.B. [det. G.O.A.,] (RNG!). 1994 ONWARDS SO80 - S0O816015 ornamental lake, Woodchester Park, 28 August 1995, R.V.L; 1998 RVL! ST78 - Wickwar — Yate area: - Goosegreen Farm, Yate Court, 5 October 1910; J.W.W. (CGE); ST709858 strontium pit, 1995, R.V.L. (RVL!); - ST710859 strontium pit, 19 August 1995, R.V.L!; ST708858 strontium pit near Yate Court, 1996 R.V.L. (RVL!). - ST717887 margin of deep quarry pool, The Cliffs, Wickwar, 19 August 1995, R.V.L! - §1T729835 newly dug ornamental ponds, Yate Golf Course, 1997, R.V.L. - §$T1T757883 pond east of Lance Coppice, 1996, R.V.L. (RVL!). ST79 - ST733967 trout pond, North Nibley, 1996, R.V.L. (RVL!). ST89 - ST899986 Cherington Lake, 23 August 1995, R.V.L! 160 R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN var. vulgaris. V.C. 6 BEFORE 1994 ST47 - Nailsea Moor, 10 September 1904, WoW. Iidet Soe SG: | (as* wvar crassicaulis det. as C. vulgaris J.G.) (BM!, E det. A.B!) (Roper 1918); ST444704 ditches; 25" January “197/55 CL E-Ge “det: J.A.M.] (BRC). ST47 - Gordano Valley: ST442735, ditch (No: 179) 3 GO Apr 9 389s NES See E.J.M.; ST443735 - ST442734, ditch (No. 186), 6 April 1989, N.F.S., S.W., E.J.M. - §T431731 ditch (No. 137), Walton Moor, 5 m altitude, 4 March 1989, N.ES., EM (SEB): - Weston Moor: ST442735 ditch (No. 7) 16" April 19895 NEES See: ST442737-ST443737, (ditch No. 148), 6 April 1989, NES S:W. (SLBI): ST442738 ditch (No. 167), 6 April 1989, N.F.S.; ST438734 peaty ditch (No. 171), 5 m altitude, 5 March 1989, N.F.S. (SLBI); ST443736 peaty ditch (No. 180), 5 m altitude, 5 March 1989, N.F.S. (SLBD); ST443737 ditch (No., 147), 6 April 1989, N.E.S., S.W. - Portbury, 24 October 1902, C.B. [det. A.B.] (BM); ditch, Portbury Marsh, October 1907, C.B. (BM). ST56 - pond between Bedminster and Whitchurch, 1882 (White 1887; Murray 1896; White 1912). V.C. 34 BEFORE 1994 ST78 - Goosegreen Farm, Yate, C.B. (White 1912; Riddelsdell et al. 1948). ST79 - ST774948 pond, Cotswold Edge Golf Club, Wotton-under-Edge, 21 January 1985, S:Greidet.J-A-Ma)) (BRE): 1994 ONWARDS SO80 - SO816015 ornamental lake, Woodchester Park, 28 August 1995, R.V.L; 1998 RVL! ST78 - ST710859 strontium pit, 19 August 1995s Revels! - §1T729835 newly dug ornamental ponds, Yate Golf Course, 1997, R.V.L. - S$T757883 pond east of Lance Coppice; 19967 RIVE. (RVE}): ST79 - ST733967 trout pond, North Nibley, 1996, R.V.L. (RVL!). ST89 - ST899986 Cherington Lake, 23 August 1995, R.V.L! NITELLOPSIS HY. Nitellopsis obtusa (Desv.) J.Groves Starry stonewort Red List - Vulnerable. A large and apparently healthy population of this species was located in a flooded gravel pit on the Frampton Pools SSSI (v.c. 34) in 1995. In 2003, populations were found in Chew Valley Lake and Blagdon Lake (both in v.c. 6), in 2004 a population was found just outside the region, in the Cotswold Water Park and in late 2004 another population was found in a flooded gravel pit in Lincolnshire. These comprise the only known sites in Britain apart from the Norfolk Broads. These discoveries over quite a short period suggest that this species may be a new arrival and be spreading. However, N. obtusa tends to be limited to fairly deep water and is therefore only found by people sampling beyond the margins who are aware of the diagnostic features. It is therefore also possible that it has been overlooked. V.C. 6 1994 ONWARD ST55 - ST515596 BlasdoniiakeweS September 2003, B.G. et al. [det. N.F.S.] (BM!). - §$T569597 Chew Valley Lake, 20 September 2003, B.G. et al. [det. N.F.S.] (BM!). V.C. 34 1994 ONWARDS SO70 - SO753076 Frampton Pools, probably this species, recorded as Nitella flexilis, 1993 C.W., A.H. (English Nature 1994); SO753076, September. 199ssaiharee (RVL!); 9 January 1996 R.V.L. (RVL!). NITELLA C. AGARDH. Nitella mucronata A. Braun (Miguel) Pointed stonewort. N. mucronata has been recorded from several sites in the region but populations have been found at only two since 1994. It is likely that all of the records refer to var. gracillima. The type locality for this variety was in a pool created by strontium extraction near Wickwar (variously “In a small pond nearly six feet deep of clear land water” (Moore & Bailey 1985), “Wickwar area’, “pond near Wickwar’ “small deep pond near Rangeworthy” by I. M. Roper). Ponds in the area which probably provided the CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET 161 type specimen were destroyed for development in late 1996. This species is nationally scarce but seems to have increased in recent years, particularly in renovated canals and it is possible that the var. gracillima is not native (Stewart 2004). V.C. 6 BEFORE 1994 ST43 - ST463394 ditch, Street Heath, 8 June 1975 P.M.W. [det. J.A.M.] (BRC); August 1989 C.F. and J.G.K [det. J.A.M.] (BRC). V.C. 34 BEFORE 1994 ST78 - ST708855 [north of the Leech Pool], Wickwar area, 1984 A.J.B. [det. J.A.M.] (BM). Nitella mucronata (A. Braun) Miq. gracillima J. Groves and Bull.-Webst. V.C. 34 BEFORE 1994 ST69 - Tortworth Lake, Tortworth Estate, 11 February 1990 N.F.S., M.A.R.K., C.K.; ST6991, 1990 RUD! (BRC); ST6992, 1990 RUD! (BRC). ST78 - Wickwar — Yate area: 11 July 1917, I.M.R. [det. G.O.A.] (BM!); 4 June 1917, I.M.R. [det. G.O.A.] (BM!); August 1918, I.M.R. [det. G.O.A.] (BM!). - small deep pond near Rangeworthy, 30 April 1917, I.M.R. [det. G.O.A.] (BM!, E!) (White 1919; Allen 1950; Moore & Bailey 1985); 4 June 1917, I.M.R. [det. G.O.A.] (BM!); pond (old strontium pit) E. of Rangeworthy, 16 May 1917, J.W.W. [det. J.G.] (BM!); pool (old strontia pit) east of Rangeworthy, 22 March 1918, J.W.W. [det. J.G.] (BM!) (Riddelsdell et al. 1948; Moore & Bailey 1986); strontia pit between Yate Court and Leech Pool Farm, Yate, 15 December 1984, A.J.B (Bailey & Moore 1985). Var. 1994 ONWARDS SO70 - SO783062 (disused) Thames and Severn Canal near Eastington, 1997, R.V.L. (RVL!). SO80 - SO786031 (disused) Thames and Severn Canal, Brimscombe, 1997, R.V.L. Nitella opaca Dark stonewort Over the last few decades N. flexilis (smooth stonewort) and N. opaca were united under one species. Recently N. opaca has been reinstated as a species in line with the accepted view elsewhere in Europe (Bryant et al. 2002). As fertile material is needed to separate the two species, there are several records which cannot be attributed to either species. In this account, records for which determination to species has not been possible, have been listed here (as WN. flexilis agg.) under N. opaca, because N. flexilis s.s. has never been confirmed from either county. All records are from slow moving or still water in medium to small ponds or from canals. Elsewhere N. opaca is frequent, particularly in western Britain, while N. flexilis is nationally scarce and is scattered in lowland areas. N. opaca appears to have declined significantly in the region, with records since 1994 only from pools in the Forest of Dean, a short length of the Severn and Thames Canal, the Gordano Valley and a marl pool at Ruskin Mill near Nailsworth. One of the reasons for the lack of recent records from sites in the Wickwar — Yate area is certainly the loss of the less basic commons overlying the coal measures, near Chipping Sodbury and Yate to urban development. The species is unlikely to be found again in this area. A small number of water bodies have been and continue to be, created in association with quarrying and with housing development and it is possible that it will reappear in one of these. V.C. 6 BEFORE 1994 ST47 - ST453726 ditch, Walton Moor, 1989 S.W. det. J.A.B. ST55 - Blagdon Lake, 5 September 1933 H.S.T. (BM). ST57 - ST558731 Bristol University Botanic Gardens, 7 May 1984 A.L. [det. J.A.M.] (BM). | ST76 - In the canal at Bath, DH.G (Babington 1834; White 1887; Murray 1896). - 1918 C.I.S. (White 1920). 1994 ONWARDS ST47_ - Gordano Valley: - Walton Moor: ST435726 ditch No. 55 [det J.A.B.]; ST436726 ditch No. 56 [det J.A.B.]; ST436727 ditch No. 57, 29 June 1989, S.W., O.M. [det J-A.B.] V.C. 34 BEFORE 1994 SO61 - SO609125 N. flexilis agg. pond with slow stream entering and leaving, Cannop, 9 November 1983, M.V.M. [det. J.A.M.] 162 R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN (Bailey & Moore 1985, Moore & Bailey 1986). ST59 - pond, Tidenham Chase, 7 July 1898 J.H.M. (BM) (Riddelsdell et al. 1948). ST69 - pond, Berkeley, 1868 J.W.W. [det. H. & J.G.] (White 1912) probably the same record listed as “pond, Berkeley Canal’, 1868 (as Chara flexilis) G.A.O.B. by Riddelsdell et al. (1948). ST78 - Wickwar — Yate area: - Yate Common: 30 April 1910, J.W.W. (CGE!); three ponds, C.B. [det. J.W.W.] (CGE) (White 1912); south of Hall End, June 1917, IL.M.R. (LSR) (White 1919); pond, 22 March 1918 J.W.W. [det. H. & J.G.] (BM!) (Barton 1919; Riddelsdell et al. 1948). - pond, Little Sodbury End, G.W.G. (Sandwith & Sandwith Moore & Bailey 1986). 1953 1954, 1994 ONWARDS SO61 - N. flexilis agg. Woorgreens Lake, northeast of the Speech House, 1993, C.W., A.H. (English Nature 1994); 1997, R.V.L.; N. flexilis agg. SO631126 pond near Fox’s Bridge, near Woorgreens, 1997, R.V.L.; N. flexilis agg. SO630128, 31 August 1998, RVI: UN, flexilis’ Vacs. SOosti27, Woorgreens Lake, 31 August 1998, R.V.L. - N. flexilis agg. SO639134 Crabtree Colliery, Cinderford, 1997, R.V.L. SO90 - §$0967008 WN. flexilis agg., Thames and Severn Canal near Coates, 7 July 1977, R.H.B., J.A.M. [det. J.A.M.] (BM) (Moore & Bailey 1986); SO0967004 abundant in (disused) Severn and Thames Canal near Coates, 1997, R.V.L. (RVL!). ST89 - N. flexilis agg., Marl pool, Ruskin Mill, 1996, J.J.D! Nitella translucens (Persoon) Translucent Stonewort Agardh. N. translucens has only been recorded from a single site in the region and there, apparently only twice. It tends to occur in the deeper parts of water bodies, rarely being visible from the margins. As such, it is possible that it has been overlooked and it may be under-recorded. Conversely, this species is very much in decline in southern England and has become extinct in several counties and may well be extinct in the region. There are few areas in v.c. 34 which are likely to be suitable, but any ponds or lakes in peaty parts of the Somerset Levels could be suitable. V.C. 6 BEFORE 1994 ST43 - Ashcott Station, Glastonbury Peat Moors, 18 June 1919 H.P. and C.I.S. (BM) (Druce 1920; White 1921); Ashcott, June 1922, H.J.G. (LSR). TOLYPELLA (A. BRAUN) A. BRAUN. Tolypella glomerata (Desv.) Leonh. Clustered Stonewort Synonym: T. nidifica var. glomerata). T. glomerata is a nationally scarce and decreasing species but is widely scattered throughout Britain. The two records from the region are from disused canals before 1985. Both sites have been surveyed since 1995 and this species now appears to be extinct in both vice counties. It continues to survive in the Cotswold Water Park in Wiltshire, a few kilometres to the east of the site at Coates. V.C. 6 BEFORE 1994 i ST33 - In one of the old clay pits, near Bridgwater, 21 August, 1899 G.R.B.-W. [det. J.G.] (BM) (Bullock-Webster 1901; Marshall 1914). - Drain near Sedgemoor Cut, May 1899, G.R.B.-W. (Bullock-Webster 1901). ST43 - Ditch, King’s Sedge Moor nr. Othery, 8 May 1899, G.R.B.-W. (BM) (Marshall 1914; 1909). ST47 - Rhine near Portbury, 1922, O.V.D. (White 1923). - §1T4573 rhyne on Clapton Moor, February 1922, C.I.S. (BM) (White 1923; Willis & Jefferies 1959). V.C. 34 BEFORE 1994 SO80 - Canal between Bowbridge and Chalford on N. side, 1907, G.A.O.B.; 1922 C.LS., N.Y.S. (Riddelsdell et al. 1948). SO90 - S0967006 Thames and Severn Canal near Coates, 7 July 1977, R.H.B., J.A.M., det. J.A.M. (BM). Tolypella intricata (Trent. Ex Roth.) Leonh. Tassel Stonewort Red List Endangered; BAP This is a rare and declining species in Britain and in recent years, it has only been recorded from a few areas, with the main British populations apparently occurring in_ the Wickwar area and on Otmoor in Oxfordshire. T. intricata is a classic species of disturbed, . ephemeral waterbodies. CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET 163 It is not immediately obvious from the list of records below, but there have been three discrete periods within which Tolypella intricata has been recorded in the Wickwar — Yate area: 1917-1918 (5 records), 1952-1955 (11 records) and 1995-2000 (17 records). The first two periods coincide with periods of increased interest in stoneworts due to recent publication of identification guides. There are no data on issues such as grazing and land management in the area for the first two periods, but we have reasonable information for the third. The early 1990s followed the end of the period of low rainfall of the early to mid- 1980s and comprised a period with good annual rainfall. Active woodland management had been carried out in Lower Woods NR resulting in poaching of many of the rides, while large numbers of cattle were grazed on the commons, to the extent that the pond to the east of Lance Coppice was usually churned to completely bare mud in the early summer. As a result of this combination of factors, from a single plant found in the pond east of Lance Coppice in 1995, eight sites had growing plants in 1997, with most supporting more than ten plants (although counts of plants have only limited meaning in relation to aquatic annual plants such as this). Since then, a decline in grazing pressure due to a combination of the foot and mouth disease outbreak and falling profits for dairy farming, combined with no significant poaching on the rides in Lower Woods NR, has led to vegetation succession in most of the ponds that previously supported T. intricata. Surveys in 2002 and 2004 found plants at only five sites in 2002, two of which were new for the species (Williams et al. 2003) and only one plant at one site in 2004. Outside the Wickwar — Yate area, the prognosis for 7. intricata is not good. The Hereford and Gloucester Canal was dry and largely full of earth in 1997, although parts were under restoration and the plant may re- appear. There is no sign of the pond near Pitney Wood, although no thorough survey of the area has been carried out. The ditch in which it was recorded in the Gordano Valley has been searched a number of times in recent years and no plants have been found, possibly because the ditch is becoming overgrown. In all these sites, except possibly at Pitney Wood, there is no reason to suppose that the species has disappeared. However without frequent visits we are unlikely to know if and when it does reappear. Similarly, without frequent survey of all sites from which the species has been recorded we are unlikely to gain a better understanding of the dynamics of populations of this species and consequently decisions on the need for management will always be somewhat speculative. The dramatic decline in the number of sites supporting plants certainly appears to be worrying, but we still have insufficient data to know whether the best course is to intervene, with the risk of creating a more or less a permanent obligation to management, or whether it is better to do nothing in the expectation that it normally undergoes a “boom and bust’ cycle and that populations will recover naturally. V.C. 6 BEFORE 1994 ST42 - pond outside Pitney Wood, nr. Langport, 21 March 1907, E.S.M. [det. G.O.A.] (BM!, E!, NMW!) (Druce 1919); pond on northern edge of Pitney Wood, 27 April 1907, E.S.M. [det. J.G.] (BM!, E!, NMW!, OXF!) (Riddelsdell 1908, Groves and Bullock-Webster 1924, Marshall 1908 — as “pond outside Pitney Wood”). ST47_ - near Bristol 1917, C.I.S. [det. G.O.A.] (BM) (Druce 1919) (possibly refers to Weston Moor). - $T441736 recently cleared peaty ditch (No. 177), Weston Moor, Gordano Valley, 6 April 1989, N.F.S. (Willis 1989) (SLBD; ST441736 ditch No. 177, 6 June 1989, N.E.S., S.W., E.J.M. ST76 - canal at Bath 1918, C.I.S. (White 1920). 1994 ONWARDS ST44 - Tealham Moor: ST40794509 ditch No. 11, 9 May 2003, N.F.S. V.C. 34 BEFORE 1994 SO72_ - ditches near Newent Canal, (as Chara vulgaris) G.A.O.B. (Sandwith 1917; Groves & Bullock-Webster 1920; Riddelsdell et al. 1948). ST78 - Wickwar — Yate area: - Yate area: In a small muddy pool (a strontium pit) north of Yate Church 25 May 1917 C.LS. [det. G.O.A.] (BM labelled “Yate Common, below Yate Rocks’) (White 1919; Sandwith 1917; Druce 1918); Different pond near Yate 1918, C.LS. [det. G.O.A.] (BM) (White 1920; Riddelsdell et al. 1948; Moore & Bailey 1986); Yate, 8 November 1918, C.IS. [det. G.O.A.] (BM) 164 R. V. LANSDOWN, N. F. STEWART, C. KITCHEN AND M. A. R. KITCHEN (White 1920; Moore & Bailey 1986); Old strontium pit below Yate Rocks 1917, (Riddelsdell et al. 1948 “in Hb. Sandwith’’); Old disused strontium digging in the neighbourhood of Yate 25 May 1917, C.LS. [det. J.G.] (Sandwith 1917; Druce 1918, Riddelsdell et al. 1948). 1994 ONWARDS ST78 - Wickwar — Yate area: - Little Sodbury and Horton: Pond, Little Sodbury End, Sodbury Common 1954, G.W.G. (Sandwith & Sandwith 1955; Moore & Bailey 1986); Sodbury Common 1954, G.W.G. (Moore & Bailey 1986); Horton 1953 G.W.G. [det. G.O.A.] (Sandwith & Sandwith 1954, Moore & Bailey 1986); Pond nr. Vinney’s Lane, Horton 1952 (Sandwith & Sandwith 1953): 1953 G.W.G. [det. G.O.A.] (Sandwith & Sandwith 1954); 1954 G.W.G. _ [det. G.O.A.] (Sandwith & Sandwith 1954; Wallace 1955; Moore & Bailey 1986). ST743851 pond nr. Vinney’s Lane, Horton, 22 February 1999, R.V.L. (Stewart & Lansdown 1999b); 5 June 1999, R.V.L., N.F.S.; 2000 R.V.L.; ST 761871 Poached, grazed and partly shaded pond on eastern edge of Hawkesbury Common, 2002, R.V.L. (Williams et al. 2003); pond, Hawkesbury Common, 24 February 2004, N.E.S., R.V.L. - Inglestone Common: Ponds, 1953, G.W.G. [det. GO.A.] (Sandwith & Sandwith 1954, Moore & Bailey 1986); two ponds 1954, G.W.G. [det. G.O.A.] (Sandwith & Sandwith 1954, Wallace 1955, Moore & Bailey 1986); 6 May 1955, H.J.M.B. (RNG!); ST758884 pond east of Lance Coppice, 21 December 1996, R.V.L. (Stewart & Lansdown 1999); 4 March 1997, RONG: 29° March 1997 ROVEE, (Stewart & Lansdown 1999b) (RVL!); 14 December 1997, R.V.L. (Stewart & Lansdown 1999); 1998 R.V.L.; 5 June 1999 N.F.S., R.V.L.; 24 February 2004, N.E.S., R.V.L.; ST751884 pond north of Spoil Coppice, Inglestone Common, 21 December 1996, R.V.L. (Stewart & Lansdown 1999b). - Lower Woods NR: ST748883 Pond, track to Lower Woods Lodge, April 1985, A.J.B. [det J.A.M.] (BM); ST748883 pond, track to Lower Woods Lodge, Lower Woods NR, Upper Wetmore, 29 March 1997, R.V.L. (Stewart & Lansdown 1999b); ST743864 track, Stonybridge Wood, Lower Woods NR, Upper Wetmore, 19 May 1997, R.V.L. (Stewart & Lansdown 1999b); ST748881 poached pool in Green Trench, south of Spoil Coppice 1999 R.V.L.; Horse’s hoof print, Horton Great Trench, Upper Wetmore 1954 G.W.G. [det. G.W.G.] (Moore & Bailey 1986); ST743873-ST741872 series of hollows in Horton Great Trench, Lower Woods NR, Upper Wetmore, 21 December 1997, R.V.L. (Stewart & Lansdown 1999b); 27 February 1999, R.V.L. (Stewart & Lansdown 1999b). - Assley Common: ST759894 pond east of Withymore Wood, nr. Inglestone, 10 April 1997, R.V.L. (Stewart & Lansdown 1999b). - ST736895 pond east of Cherryrock Farm, nr. Inglestone, 9 April 1997, R.V.L. (Stewart & Lansdown 1999b) (RVL!). - ST718888 margin of deep quarry pool north of Wickwar, 19 May 1997, R.V.L (Stewart & Lansdown 1999b) (RVL!). Tolypella nidifica (O. Mill.) Leonh. Bird’s-nest stonewort This species was “Alleged to have been detected in old river bed at Sharpness (C.N.F.C. Ill, 1863, 145 [presumably indicates that it was published in the Proceedings of the Cotteswold Naturalists Field Club]), but doubtless in error’ (Riddelsdell et al. 1948). The original record was probably given as Chara nidifica (although even this is a little late for use of this name). C. nidifica auct brit. has tended to be equated with Tolypella glomerata, but could also have referred to Nitella opaca or N. flexilis. Thus, whilst Riddelsdell et al. (1948) robustly dismisses this record and it is very unlikely to have been this species, it could have been a Tolypella species, particularly T. glomerata or T. intricata and it may be worth searching this area for suitable habitat for both taxa. Tolypella prolifera (Ziz. ex A. Braun) Leonh. Great Tassel Stonewort Red List — Endangered; BAP Before 1994, T. prolifera had been reported from two sites in the region. Survey by the Somerset Rare Plants Group at Southlake Moor resulted in the discovery of an apparently new, healthy population of 7. prolifera. In contrast to all other British sites for this species, where CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET 165 populations are normally restricted to one or two ditches, at both sites in the Somerset Levels it has been record from ten ditches. There have apparently been no recent surveys of the gravel pits in the Walham area by botanists familiar with this species. Not only do some of these pits support uncommon aquatic plants, such as Ceratophyllum submersum, but there are abundant ditches and small ponds in the area that may be suitable for this species and which merit survey. V.C. 6 1994 ONWARDS. Siisse—s36393048; “ditch No. 53; ST36863056, ditch No. 60E, 26 June 1998, Reverses Po LW. (leach et. al. 1999); 18 July 2000, N.F.S., K.P. (Williams & Stewart 2002); ST36243055, ditch No. 1; ST36313057, ditch No. 3; ST36773056, ditch No. 30; ST36843057, ditch No. 31; ST36903058, ditch No. 32; ST36863056, ditch No. 60E; ST36913057, ditch No. 61; ST36933040, ditch No. 97: ST36983045, ditch No. 98; ST37043045, ditch No. 99N, 20 July 2000, N.F.S., K.P.; 26 July 2000, N.E.S., K.P., P.W. (Williams & Stewart 2002) la) August 2000, P.W., J.B. (Williams & Stewart 2002); ST36843057, ditch No. 31; ST36333026, ditch No. 85; ST36373030, ditch No. 88; ST36933040, ditch No. 97; ST36983045, ditch No. 98, 26 July 2000, N.F.S., P.W., K.P.; 4 October 2001, N.F.S. (Williams & Stewart 2002); ST36723054, ditch No. 78 N_ end; ST36813052, ditch No. 79; ST36933040, ditch No. 97; ST36783030, ditch No. 147, 23 October 2001, N.F.S. (Williams & Stewart 2002); ST37413027, ditch No. 120; ST53 - ST5236 in a small ditch surrounded by frogbit, September 1986, J.G.K. Butt Moor (Leach et. al. 1999); ST52563606 ditch No. 2, 26 July 2000, N.F.S., P.W. (Williams & Stewart 2002); ST52563606 ditch No. 2, 15 August 2000, P.W., J.Bi. (Williams & Stewart 2002); 23 October 2001, N.ES. (Williams & Stewart 2002); ST52623609 ditch No. 3; ST52613600 ditch No. 4, 10 October 2002, N.F.S.; ST52503611 ditch NOM le Silis2563606) ditch’ Not) 2: ST52503585 ditch No. 8; ST52223608 ditch No. 11; ST52173610 ditch No. 12; ST52213615 ditch No. 13, 23 July 2003, N.F.S., E.J.M. et al.; ST52463583 ditch No. 9; ST52663563 ditch No. 10; ST52223608 ditch No. 11; ST52263640 ditch No. 14, Butt Moor 17 September 2003, N.F.S. V.C. 34 BEFORE 1994 SO82 - Walham, 4 July 1939, J.F.G.C., N.D.S [det. G.O.A.] (BM). ACKNOWLEDGMENTS The authors would like to thank Chris Preston of the Centre for Ecology and Hydrology for providing information held on the B.R.C. database and Charlotte Pagendam and Mike Wilkinson of English Nature, Three Counties Team for providing data from the open waters survey and for background information on Frampton Pools S.S.S.I. In addition, R. V. Lansdown would like to thank Barbara Costello for access to the Biological Science Library of the University of Bristol, Peter Martin for access to his copy of Dandy (1969) and Dr. A. ST36903034, ditch No. 137E, 19 R. Lansdown for information on the uses of September 2003, N.F.S. strontium. REFERENCES ALLEN, G. O. (1950). Nitella mucronata Mig. var. gracillima Groves and Bull.-Webst. in Warwickshire. Watsonia 1: 364-365. BABINGTON, C. C. (1834). Flora Bathonensis. London. BAILEY, R. H. & Moore, J. A. (1985). Gloucestershire charophyte records 3. Gloucestershire Naturalists Society Journal 36: 80-83. BARTON, W. C. 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KITCHEN THE NUMBER OF 10 KILOMETRE SQUARES WITH RECORDS OF EACH TAXON Taxon Chara aculeolata Chara aspera Chara contraria Chara globularis Chara hispida Chara virgata Chara vulgaris var. crassicaulis var. longibracteata var. papillata var. vulgaris Nitellopsis obtusa Nitella mucronata var. gracillima Nitella opaca Nitella translucens Tolypella glomerata Tolypella intricata Tolypella nidifica Tolypella prolifera Total IUCN No. 10 km squares v.c. 6 No. 10 km squares v.c. 34 pre-1994 post-1994 pre-1994 post-1994 NS [1] 1 [1] 2 2 4 7 + 6 1 6 1 yy 1 3 1 5) 4 16 8 18 9 1 2 Z 4 4 7 3 6 4 3 Z 4 VU, BAP Z 1 NS 1 1 2 2 + 1 5) 3 1 NS 3 yy EN, BAP 3 1 2 1 EN, BAP [1] EN, BAP 2 1 59[2] 28 56[1] 38 The numbers given represent the total number of 10 kilometre squares from which each taxon has been recorded in each period. Square brackets indicate doubtful records KEY: EN - Endangered VU - Vulnerable NS - Nationally Scarce - listed as a Priority Species in the UK Government Biodiversity Action Plan BAP APPENDIX: INDIVIDUALS REFERRED TO IN THE TEXT A.B. — A. Bennet A.H. — A. Harrison A.J.B.— A.J. Byfield A.J.D. — A. J Dodd A.K. — A. Kelham A.L. — A. Leitch A.Le. — A. Leipner A.O. — A. Orange A.W. — A. Washbrook B.A.G. —B. A. Grindey B.G. — B. Goldsmith C.B. — C. Bucknall C.C.B.—C. C. Babington C.F. — C. Furness C.LS. —C. I. Sandwith C.K. — C. Kitchen C.T. —C. Trapnell C.W. —C. Walker D.C. — D. Callaghan D.F. — D. Fry D.F.L. — D. F. Leney D.H. — D. Heaver D.Ha. — Dr. Hassé D.H.G. — Dr. H. Gibbes D.McC. — D. McClintock D.M.-S. — D. Munro-Smith D.N.A. —D. A. Allen DR. — D. Reid E.G. — E. Green E.H.R. — E. H. Read E.J.M. — E. J. McDonnell E.S.M. — E. S Marshall F.G. — F. Gibbons F.K.M. — F. K. Makins G.A.O.B.—G. A. O. St. Brody G.C.D. — G. C. Druce G.0.A —G. O. Allen G.R.B.-W. — G. R. Bullock- Webster . G.W.G. — G. W. Garlick H. — Hanson H.G. — H. Groves H.J.M.B.— H. J. M. Bowen H.J.R. — H. J. Riddelsdell H.K.A.S. — H. K. Airy Shaw H.P. — H. Perrycote H.P.R. — H. P. Reader H.S.T. — H. S. Thompson I.F.G. — I. F. Gravestock I.M.R. — I. M. Roper J.A.B.—J. A. Bryant (née Moore) J.A.M. — J. A. Moore (now Bryant) J.Bi. —J. Biggs J.D.H. — J. D. Hooker J.E.L. — J. E. Lousley J.G. — J. Groves J.G.K. —J. G. Keylock J.H.M. — J. H. Morgan J.J.D. —J. J. Day J.N. — J. Norton CHARACEAE IN W. GLOUCESTERSHIRE AND N. SOMERSET 169 J.P.M.B. —J. P. M. Brenan J.v.R. —J. van Raam J.W.W. —J. W. White K.E.L. — K. E. Ludbrook K.P. — K. Pollock L.B. — Rev. L. Bloomfield L.W. —L. White M.A.P. — M. A. Palmer M.A.R.K. — M. A. R. Kitchen M.D.C. — M. D. Chitty M.J.A. —M. J. Allen M.R.H. — M. R. Hughes M.S. — M. Scruby M.V.M. — M. V. Marsden N.F.S. — N. F. Stewart N.Y.S.—N. Y. Sandwith O.M. — O. Mountfield O.V.D. — O. V. Darbishire P.M. — P. Millman P.M.W. — P. M. Wade P.W. — P. Williams R.H.B. — R. H. Bailey R.P.M. — Rev. R. P. Murray R.S.C. — R. S. Cropper R.U.D. — Mrs. Ruddle R.V.L. — R. V. Lansdown R.W. —R. Wilder S.C.H. — S. C. Holland S.H.B.—S.H. Bishop S.J.L. — S. J. Leach S.P.P. —S. P. Phillips S.W. —S. Wilson T.C. — T. Clarke V.H. — V. Hack W.B.W. — W. B. Waterfall W.J. — W. Joshua W.L. — W. Lutley W.R.M. — W. R. McNaib W.W. — W. Watson Watsonia 26: 171—178 (2006) 17] J. H. Penson: mystery man and mystery record M. J. Y. FOLEY* Faraday Building, Department of Biological Sciences, University of Lancaster, Lancaster LAI 4YA ABSTRACT J. H. Penson was a little-known British botanist who made several important contributions to plant re- cording. A biographical note, together with in- formation relating mainly to his activities in Britain but also on the North American continent, is given. KEYWORDS: Carex capitata, Newfoundland, plant records, Scotland, World War 1. INTRODUCTION In late 2004, the chanced sight of an undated, unaddressed letter written by the late Scottish botanist Robert Mackechnie (1902-1978) led me to a fascinating series of enquiries. In this letter (in litt. to A. C. Jermy), when discussing J. W. Heslop Harrison’s finds on the Isle of Rhum in the 1940s, Mackechnie stated “My recollection of Carex capitata is by now rather vague, but I have the impression that it resembled J. H. Penson’s Ben Hope [Scotland] plant more closely than it did material which I saw in Lapland.....”. J. H. Penson’s Ben Hope plant indeed! Carex capitata (other than Heslop Harrison’s dubious 1940s records from the islands of Rhum and South Uist) has never been confirmed from the British Isles although, as an Arctic-montane plant, it is quite frequent in Scandinavia and Iceland and, conceivably, could also occur here, especially in northern Scotland. So Penson’s record for Ben Hope could be of particular significance. To follow this up, therefore, one needed to locate the published source (if any) of such an important record and, if a specimen had been collected, to locate and examine the voucher. This, however, was where the difficulties began. Who was J. H. Penson? Wide-ranging enquiries amongst botanists, especially Scottish, who might have known, or heard of him, led virtually nowhere. A few people vaguely recalled the name but knew little about him and only one person re- membered once meeting someone referred to as *e-mail: m.foley @lancaster.ac.uk “old Penson”. A request for information in current botanical publications also met with a similar minimal response, whilst a search through past British botanical literature for any records he might have provided mostly drew a blank. His name is mentioned in_ the introductory section of The Changing Flora of Glasgow (Dickson et al. 2000) along with a record for Cirsium heterophyllum and there are other occasional records in a few issues of the The Glasgow Naturalist but, in neither, is biographical information forthcoming. In addition, and most surprisingly, there appears to have been no published obituary of Penson (assuming him to be no longer alive). An Internet search, however, revealed the existence within the first half of the twentieth century of someone with that surname and initials apparently residing in the U.S.A. and Newfoundland but without any definite British connection stated. The breakthrough came by chance and in rather unfortunate circumstances. During house clearance following the death of Allan McG. Stirling in 2004, a series of botanical note- books dating from 1958 onwards, together with a few miscellaneous letters and ephemera, were located; these were clearly the former possessions of J. H. Penson. From such early clues and a firmer establishment of Penson’s past connection with the North American continent, it has been possible to provide what is still a rather incomplete summary of some important and interesting facts about this almost unknown, elusive British botanist. A BRIEF BIOGRAPHY Hubert Penson was born on January 21, 1893, at Hornsey, London and died on June 1, 1979, in the Southern General Hospital, Glasgow. He was the son of Arthur A. Penson, a dairy manager of Cirencester. In 1929 he married Marjorie Doreen Crawford (deceased 1971) with whom he had one son and two daughters. In 1974 he married again, this time Ellen Mary Cumming (deceased 1991). WZ To date, details of his early life and education are lacking and it is not until his involvement in World War 1, where he served from 1916-1919, that information becomes available. In this conflict he was _ twice decorated for bravery. As Lt. Hubert Penson (promoted to that rank in 1918) of the 5th Division Signal Company of the Royal Engineers and attached to the 15th Infantry Brigade, he was awarded the Military Cross (MC) in 1918. For this, the citation recognised his “conspicuous gallantry and devotion to duty in maintaining communications which had to be constantly repaired under heavy shell fire. In addition to carrying out his signal duties he sent important information when the situation was obscure”. About a year later, serving in northern Russia, he was awarded a bar to his Military Cross. This recognised his “conspicuous gallantry and devotion to duty on August 10, 1919, having guided the Sludka- Lipovets Column for three days, and brought it to within 400 yards of the enemy’s position unobserved”. On the same day, “the column, having to withdraw from Lipovets owing to shortage of ammunition, he [Penson] guided it out under heavy fire’. He was also mentioned in military despatches. After that the trail goes cold until 1923 at which time he wrote an article entitled ‘iransiations, And) thew Necdin som Guidance’ (Penson 1923). In this, he stated: “Outwardly men differ widely — in language, in appearance. Inwardly it is the similarity that is remarkable...... Translators of the Russian writers — those of Tolstoy, for example — have to give the world this world-wide appeal”. About the same time he moved into Government finance and was still living in the London area as evidenced by _ recently- discovered letters addressed to him, one of which indicated his residence to be in Well Walk, Hampstead. In November 1928, he was appointed Assistant Secretary to the Com- mission on Museums and Galleries and two years later wrote an article in the Journal of Royal Institute of International Affairs under the title “The Financing of Russian industry”. His rise in Government circles must now have been rapid for in 1931 he became a member of the Newfoundland Royal Com- mission and subsequently was appointed there as Commissioner for Finance. Around this time he appears to have spent periods on both sides of the Atlantic and the inference gained from a letter to him dated March 1941 suggests that he M. J. Y. FOLEY retained his house in Hampstead, London, until almost 1940. Despite this, from the mid-1930s, his career was apparently spent mainly on the North American continent. In1937, he lectured on topics of contemporary interest at Memorial University College (St John’s, Newfoundland) and in July 1938 issued a report stating Newfoundland’s revenues to be the greatest in the country’s history. This, no doubt, was a reflection on his astute management! In the following year he advised on slum clearance in St John’s but found the project to be financially impractical. In July 1940, as Commissioner of Finance, he stated that, for the remainder of the war, Newfoundland would no longer ask Britain for financial assistance and a year later announced a $700,000 budgetary surplus. In 1940 he was sent to St Pierre-et-Miquelon (the French enclave in Newfoundland) to meet with the Administrator in order to assess the attitude to the Nazi axis powers. He found this to be distinctly anti-Axis and, in addition, discovered that St Pierre would offer few useful facilities as an enemy military base. Shortly afterwards, his work in Newfoundland was complete and in February 1941 he was replaced as a member of the Commission of Government in New- foundland. In 1942, for his past services he was awarded the Cross of St Michael and St George (CMG ). There is then a three-year gap until 1944 when he became Secretary-General to the British Supply Mission in Washington. Whether he remained on the western side of the Atlantic all this time is unclear, but bearing in mind the hostilities of World War 2, it is likely. Following the war, he was appointed Attaché to the British Embassy in Washington and remained in that post until his retirement, meanwhile co-authoring a book on the history of Poland for the years 1697-1935 (Reddaway et al. 1951). In the New Year Honours List for 1953, he received a further award, the Order of the Bath (CB), for his services at the Washington Embassy. From then he appears he to have been in semi-retirement, moving back to Britain and living in Nithsdale Road, Glasgow. From this base he botanised over much of the north of Britain and, especially, Scotland. Surprisingly, on his second marriage in 1974 at the age of 81, his occupation was described as “Sales Representative” so he was perhaps still partly employed, even at that age. His rather flimsy, but well-kept botanical note-books conclude abruptly with an entry for a visit to Upper J. H. PENSON 173 Dallochy, Moray, on July 21, 1978. Possibly this was the result of the onset of a sudden illness, as he died ten months later. BOTANICAL EXPLOITS To date, the first indication of Penson’s interest in botany is a pressed fragment of Linnaea borealis. This, now in E, is marked “N[orth] Russia, 1919”, and was presumably collected at the time of his military exploits there with the Sludka-Lipovets Column. Other than this, his known botanical work falls into three phases: (a) early activities in Britain up to the mid- 1930s for which there is now only scant evidence; (b) investigations of the New- foundland (now Canada) and later Washington floras from that date until the mid-1950s. Information on this period is currently un- processed and held at the Université de Montréal, herbier Marie-Victorin (MT); (c) his semi- or post-retirement period in the British Isles, mostly Scotland, from 1958 when he made several new or ffirst post-1930 “Atlas” (Perring & Walters 1962) records for several vice-counties. A. THE PERIOD PRIOR TO THE MID-1930S: It was only following the discovery of a file containing letters sent to Penson, that anything at all could be said about this time, and even this information is very sparse. The letters only date back to 1933 but, other than the Linnaea Specimen mentioned above, are the first indication of Penson’s botanical interest. In them are discussions (with Clarice Vurian) regarding Potentilla fruticosa in the Welsh Borders and Spiranthes and Calamagrostis in Ireland, and an offer from Francis Druce to propose Penson for membership of the Linnean Society (which he apparently never took up). Around this time he was in discussion with A. H. G. Alston regarding Cystopteris in Teesdale and also showed an interest in the sugar limestone formations of the area. He also corresponded with F. Godwin regarding vari- ous orchids and Pulsatilla vulgaris. B. NEWFOUNDLAND AND WASHINGTON c.1935 TO THE MID-1950S: The small amount of information so far located on Penson’s records and collections from the North American continent date from the last few years of this period. He was however, active over much of the full period and, prior to his return to Britain, donated his collection of Newfoundland plants to the Université de Montréal, herbier Marie-Victorin (MT). There are extant letters (both in MT and now in E) in which this gift is discussed with Ernest Rouleau, the curator at that time. In one, dated December 10, 1951 (now in E) and addressed to Penson at the British Embassy, Washington, Rouleau states that “especially appreciated would be Graminae, Cyperaceae, Salicaceae, Caryophyllaceae, Draba, Antennaria, Senecio, Epilobium. Also, Dryas drummondii, Myosotis arvensis and Gnaphthalium supinum” and that his [Penson’s] “species are considered to be historically important and should be in a Canadian herbarium’. At present, in MT, there are two copies of a 72 page unpublished manuscript by Penson under the title ‘List of Newfoundland Plants’, one copy of which is subsequently annotated by him. There are also additional documents, including a plant list dealing with questions which he had put to Rouleau together with the latter’s answers. In addition, his collecting expeditions and explorations are mentioned in the Atlas of the vascular plants of the island of Newfoundland and of the islands of Saint-Pierre-et-Miquelon (Rouleau & Lamoureax 1992). As far as the specimens are concerned, these are still being incorporated at MT, but within the database produced to date, there are already over 250 collections with many more to be added (pers. comm. S. Hay). In addition, other specimens are held at the U.S. National Herbarium at the Smithsonian Institution, Washington (US) and at the New York Botanical Garden (NY). A note in the journal Rhodora (Smith 1953) refers to a Penson record for Borrichia frutescens, this confirming the presence of the plant in the District of Columbia whilst in a letter (now in E) the Curator of Ferns (US) acknowledges his gift of material of Ophioglossum engelmanni. So, it is to the specimens and data held at the above institutions, especially that at MT, that further investigations of his time over there should be addressed. C. THE BRITISH ISLES (MAINLY SCOTLAND) 1955-1978: Apart from a recently discovered herbarium specimen of Bartsia alpina collected on Ben Lawers in August.1955 (EK), the earliest of his records (and notebooks) so far traced date from 1958; this was some time after he moved to Glasgow on his return from the U.S.A. For the first few years of this period he botanised mainly in the central Scottish belt. As his interest and experience of Scottish plants Ne MINS FOLEY widened he ventured further afield, especially to the north and north-west, to Perthshire, to the Clova hills, the Cairngorms and the far north, as well as occasionally into southern Scotland, northern England and even to Ireland. In much of this time he botanised almost the whole year round. To West Sutherland (v.c. 108) alone he made over thirty separate visits, and to Ben Hope, apparently a favourite of his, at least ten. During the 1960s, like many before him, he sought out British rarities which involved visits to the Cuillin for Arabis alpina, the Storr, Skye (v.c.104) for Koenigia islandica, Cul Mhor, W. Ross (v.c. 105) for Artemisia norvegica and to Meikle Kilrannoch (v.c. 90) and Hobcarton Crag, Cumbria (v.c. 70) for Lychnis alpina. He must have been a person with a strong physical constitution since many of his trips involved walks over long distances in rough mountain terrain. Several remarkable excursions were made when he was in his mid-seventies. One, at the age of seventy-seven, was to the distant corries of Beinn a’Bhuird and the northern Slochd Mhor side of Ben Avon (v.c. 94) to search for Saxifraga cespitosa. Although failing to find it in earlier visits, he eventually recorded about 20 “cushions” there on September 24, 1970. Even when over eighty he was tackling Ben Heasgarnich, Ben Hope and Ben Lomond. Although a few of his plant identifications are rather fanciful, the majority seem reliable and he was never reluctant to attempt to identify hybrids or taxa down to subspecific and (occasionally) to varietal level. He showed a particular interest in critical genera, such as alpine and sub-alpine Hieracium and Salix and, from Ben Hope alone, recorded 21 taxa of the former and several hybrids within the latter. As a botanist, Penson appears to have been a ‘loner’, only rarely in contact with a colleague, although Robert (Bob) Mackechnie (1902-— 1978) was someone to whom he is known to have written and who also examined some of his rarer specimens. Penson’s find of Carex elongata at Loch Lomond (see Appendix 1) was probably the start of their association. The only other persons with whom he seems to have kept in touch were Allan McG. Stirling and Basil W. Ribbons. During his years in Scotland, Penson made several significant finds including first vice- county and first post-1930 “Atlas” records. Some of the more interesting of these are listed in Appendix 1. As an example of his botanical ability and his willingness to thoroughly explore individual localities, Appendix 2 lists over 160 taxa which he recorded between the years 1969 and 1975 on his favourite mountain, Ben Hope. It is hoped that these records might be of value as a basis for further recording work there. On his death, Penson’s British herbarium passed to the Nature Conservancy Council (N.C.C.), South-West Scotland Office, at Balloch around about 1980 (see letters by his widow and A. McG. Stirling (now in E)). When this organisation (N.C.C.) underwent reorganisation, the material was transferred to Scottish Natural Heritage (S.N.H.) Regional Headquarters at Clydebank (pers. comm. J. Mitchell, 2005). Unfortunately, it has not been possible so far to locate its current whereabouts. However, not all of his collections appear to have been included in the N.C.C. bequest since some of them were retained by A. McG. Stirling. These should come to light when his personal collection is fully processed and incorporated into the Edinburgh herbarium (E). An interesting sequence of records relating to a population of Hierochloe odorata date from his early time in Scotland and run through to the mid-1970s. This was at the (apparently) threatened Blythswood (v.c. 76) site by the River Cart which was first discovered in 1931 (Dickson et al. 2000). Under its earlier name (Savastana odorata L.), Penson recorded 20— 25 flowering plants there in April, 1959 followed by 12 and 64, respectively, in the following two years. In 1962 there were 18, the site having been partly damaged by refuse dumping, whilst in the following year it was entirely covered with water-borne debris and no plants were seen. In 1964 and 1965 there were also none, Penson describing the area as “obliterated by oil and rubbish”. However, following the effects of a violent storm in January 1968, much of the rubbish was washed away and 60 plants now flowered. The good news was short-lived, for the following year he described the site as “destroyed” by burnt rubbish with no sign of the plant whatsoever. However, there was another revival, and from then on, despite continued despoliation there, he continued to find the plant as two separate populations right up to his last visit there in 1976. In the early 1970s the number of flowering plants roughly fluctuated between a total of 40 and 80 with a peak of 132 in 1971. J. H. PENSON 175 Records for his last two seasons showed only 11 and 5 respectively, Penson again expressing concer regarding the deteriorating conditions nearby and the threat of development. However, despite all his apprehension, Hiero- chloe is still present at the site with two patches on the east side of the river and one on a small island to the west (pers. comm. K. Watson 2006) and these populations have been recently studied by Tait (2000). SUMMARY John Hubert Penson, hitherto virtually unknown and unremembered (at least in botanical circles), appears to have been a person of no mean talent. His exploits in World War 1 and his professional career in British Government circles, mark him as someone out of the ordinary. There is still much more to learn about Penson and especially of his botanical work on the North American continent. A summary of this may be provided later when data held at the Université de Montréal, herbier Marie-Victorin (MT) becomes available. The series of letters written to Penson, his extant field note-books, and most of the peripheral information on which this account is based, is now deposited at the herbarium/ library of the Royal Botanic Garden, Edinburgh. There, additional herbarium material is currently being incorporated which will, no doubt, contain additional examples of his collections. In consequence, more information should then be forthcoming. Finally, the point where all this started: Carex capitata: In one of the Penson note- books, the entry for July 30, 1973, states: “Ben Hope, W. Suthd., W side to 1500 ft, Carex capitata”. Unfortunately, the epithet capitata has been struck out and the comment “re- identified as C. dioica” inserted in Penson’s hand. That is probably the last word on the record unless his actual specimen comes to light and, after all, confirms his original determination. Nevertheless, the presence of C. capitata in northern Scotland remains a distinct possibility. All herbarium codes quoted above are as given in Holmgren ef al. (1990). A detailed examination of Penson’s note-books might be of interest to present-day Scottish botanical recorders. ACKNOWLEDGMENTS In compiling this account I am very grateful to the following for the information and help they have provided: I. Hedge, E. Imery, A. C. Jermy, D. McCosh, D. McKean, J. McNeill, P. McPherson, J. Mitchell, J. Robinson, A. Rutherford and K. Watson (all of Scotland). Also to T. Watkins (the Central Chancery, London) and to L. Brouillet and S. Hay (both of Montreal). REFERENCES DICKSON, J. H., MACPHERSON, P. & WATSON, K. J. (2000). The Changing Flora of Glasgow. Edinburgh University Press, Edinburgh. HOLMGREN, P. K., HOLMGREN, N. H. AND BARNETT, C. (1990). Index Herbariorum. Part 1: The Herbaria of the World. New York Botanic Garden, New York. MITCHELL, J. & STIRLING, A. MCG. (1980). Carex elongata in Scotland. The Glasgow Naturalist 20: 65-70. PENSON, J. H. (1923). Translations and the Need of Guidance, The Author 75—76. PERRING, F. H. & WALTERS, S. M. (1962). Atlas of the British Flora. Thomas Nelson, London. REDDAWAY, W. F., PENSON, J H., HALECKI, O. & DYBOSKI, R. (1951). The Cambridge History of Poland from Augustus 11 to Pilsudski (1697-1935). Cambridge University Press, Cambridge. ROULEAU, E. & LAMOUREUX, G. (1992). Atlas of the vascular plants of the island of Newfoundland and of the islands of Saint-Pierre-et-Miquelon. Fleurbec, Québec. SMITH, L. B. (1953). Borrichia frutescens from Chesapeake Bay. Rhodora 55: 58-59. STIRLING, A. McG. (1978). Common Broomrape, in Mackechnie, R., Naturalist 19: 426-427. ed. Short Notes. The Glasgow TAIT, T. N. (2000). The history, habitat and present status of Holy Grass (Hierochloe odorata (L.) P. Beauv.) at Blythswood, Renfrew District, v.c.76. The Glasgow Naturalist 23(5): 17-20. (Accepted May 2006) 176 M. J. Y. FOLEY APPENDIX 1 SOME OF PENSON’S MORE INTERESTING SCOTTISH FINDS (FOR SOME OF WHICH VOUCHERS ARE AVAILABLE) Allium carinatum: waste ground by River Clyde, Bowling, Renfrewshire (v.c. 76), DOSS) (KE) Betula pubescens: Glen Doll, Angus (v.c. 90), 25.7.1961 (a form approaching Betula nana) (E) Botrychium lunaria: an abnormal form with three fertile branches originating from the sterile base of the frond; railway bank, near Cononish River, Tyndrum, Mid Perth (v.c. 88), 4.vi.1974 (E) Carex buxbaumii: near Arisaig, Westerness (v.c. 97), 5.8.1964 (lacking an inflorescence) (E) Carex elongata: first recorded by Penson north of Balloch, near Loch Lomond, Dunbarton (v.c. 99) on June 16, 1967 (E). He had twice been in the same general area in the previous year but failed to record it. Three years later on June 27, 1970, he recorded “about 75 plants” flowering freely c.1400 yards north of the northern edge of Balloch Park and confined to a small area of about one acre (notebook); also same date and apparently same place in “wet woods” (E); first post- 1930 “Atlas” record for vice-county. These finds probably led to his introduction to Mackechnie and Ribbons. Further details can be found in Mitchell & Stirling (1980). Carex punctata: Rocks of Garchew, Wigtownshire (v.c. 74), 5.9.1962 (E); first post-1930 “Atlas” record for vice-county. Cicerbita alpina: Craig Maud, Clova, Angus (v.c. 90), 31.8.1960 (GL), notebook states “...only 4 plants left, poor state, no flowers, looked as if site robbed”’. Cirsium heterophyllum: Pollok, Glasgow, Renfrewshire (v.c. 76), 15.6.1959 (see notebook); also Dickson et al. (2000). Dryopteris assimilis: “mature spores were lacking”, Meall nan Tarmachan, Killin, W. Perth (v.c. 87), 9.10.1965 (The Glasgow Naturalist, xviii(9): 520 (1967)). Geranium purpureum: Largybaan, Mull of Kintyre (v.c. 101), 17.7.1968 (a dubious determination) (E). Hieracium alpinum: Beinn a’ Bhuird (v.c. 92), 3000’, 8.8.1968 (E). Hierochloe odorata: Banks of River Cart, Blythswood, Renfrewshire (v.c. 76), 24.5.1969 (E) (for more details see the account above). Lathyrus montanus var. tenuifolius: Ben Hope, W Sutherland (v.c. 108), lower slopes, WQS (2). Ledum palustre: Barochan Moss, Renfrewshire (v.c. 76), 2.9.1975 (E), a new v.c. record. Orobanche minor: Bowling, Dunbarton (v.c. 99), regularly recorded here by Penson in the years 1964-1977, a small colony with a maximum of 20 plants in 1970 (see also Stirling 1978). He attributed the Bowling plants to the taxonomically obscure var. conciliata Druce (see notebook). Orobanche rapum-genistae: Locharbriggs, near Dumfries (v.c. 72), 9 plants on 24.8.1966 (see notebook). Potentilla argentea: Balthayok, East Perthshire (v.c. 89), 7.6.1960 (E), first post-1930 “Atlas” record for vice-county. Potentilla tabernaemontani: Ferry Hills, North Queensferry, Fife (v.c. 85), 30+ plants on 24.4.1973 (see notebook); first post-1930 “Atlas” record for vice-county. Poa xjemtlandica: Ben Dorain, Argyll (v.c. 98), 1960 (GL). This is definitely an error and is possibly of Deschampsia alpina. However, the specimen is now very degraded and is not possible to be determined accurately. Despite this Penson did record Poa xjemtlandica on Lochnagar, S. Aberdeen (v.c. 92), (a known locality) on 28.8.1962. Polygonatum verticillatum: Den of Reichip, near Dunkeld, East Perth (v.c. 89), 5.8.1961 (E); first post-1930 “Atlas” record for vice- county. Ulex gallii: 3 miles south of Forsinard, East Sutherland (v.c. 107), 8 clumps in full flower, 17.10.1967; a new v.c. record. n.b. “Atlas” refers to Perring & Walters (1962) J. H. PENSON a7 APPENDIX 2 PENSON’S PLANT RECORDS FOR BEN HOPE (V.C. 108); (OCTOBER 1969-JUNE 1975) Agrostis canina subsp. montana Alchemilla filicaulis Alchemilla glomerulans Antennaria dioica (in many forms) Arctostaphylos uva-ursi Arctous alpina Athyrium alpestre Betula nana x B. pubescens Betula pubescens x B. nana Botrychium lunaria Caltha palustris subsp. minor Cardamine flexuosa Cardaminopsis petraea Carex canescens (= C. curta) Carex capitata (re-identified as C. dioica!) Carex curta x C. echinata (= C. xbiharica) Carex demissa Carex dioica (or hybrid) Carex elata Carex hostiana Carex lepidocarpa Carex livida Carex oederi Carex pallescens Carex pilulifera Carex pilulifera var. longibracteata Carex pulicaris Carex serotina Catapodium rigidum Cerastium alpinum Cherleria sedoides Cochlearia alpina Cochlearia officinalis subsp. alpina Coeloglossum viride Cornus suecica Cystopteris fragilis var. dentata Dactylorchis fuchsii subsp. hebridensis Dactylorchis fuchsii subsp. hebridensis x subsp. fuchsii Dactylorchis fuchsii subsp. o’kellyi Dactylorchis occidentalis Dactylorchis praetermissa var. pulchella Dactylorchis purpurella Dactylorchis purpurella x D. majalis Deschampsia alpina Draba (query) norvegica (later corrected to Cardaminopsis petraea) Drosera anglica Drosera intermedia Drosera rotundifolia x D. anglica Dryas octopetala Dryopteris abbreviata Eleocharis multicaulis Eleocharis quinqueflora Eleocharis uniglumis Empetrum hermaphroditum Epilobium alsinifolium Epilobium anagallidifolium Eriophorum gracile Euphrasia borealis Euphrasia brevipila var. subglandulosa Euphrasia confusa (and query) a form of E. micrantha Euphrasia eurycarpa (confirmed) Euphrasia frigida Euphrasia nemorosa vat. collina Euphrasia occidentalis var. calvescens Euphrasia rhumica Euphrasia scottica Festuca capillata Festuca ovina subsp. tenuifolia Festuca ovina vat. capillata Festuca pratensis Festuca vivipara Galium sterneri (a new station) Glyceria fluitans Glyceria xpedicellata Hieracium (sect. Cerinthoides) Hieracium anglicum Hieracium argenteum Hieracium backhousei Hieracium calenduliflorum Hieracium callistophyllum Hieracium cremnanthes Hieracium euprepes Hieracium eximium Hieracium eximium vat. tenellum Hieracium gracifolium Hieracium gracilentum Hieracium hanburyi Hieracium hypochaeroides Hieracium pseudanglicum Hieracium pseudocurvatum Hieracium schmidtit Hieracium senescens Hieracium sp. (Sect. Vulgatum) Hieracium submurorum Hieracium anglicum Holcus mollis Hyperichum pulchrum Ilex aquifolium Juncus alpinus Juncus triglumis Juniperus nana 178 M. J. Y. FOLEY Lathyrus montanus var. angustifiolius Lathyrus montanus var. tenuifolius Leontodon autumnalis Lobelia dortmanni Loiseleuria procumbens Luzula spicata Melampyrum pratense Melampyrum pratense subsp./var. montana Montia chondrosperma Myriophyllum alternifolium Nymphaea occidentalis Orchis mascula Pedicularius palustris Pinguicula lusitanica Platanthera bifolia Platanthera chlorantha Polygala serpyllifolia Polygala vulgaris Polystichum lobatum var. lonchitides Polystichum lonchitis Prunella vulgaris Pseudorchis albida Ranunculus flammula var. tenuifolius Rhinanthus borealis (and in pure form) Rhinanthus borealis x R. minor Rhinanthus minor subsp. stenophyllus Rhinanthus minor subsp. borealis Rhinanthus minor subsp. minor x_ subsp. borealis (R. xgardineri) Rhinanthus minor subsp. monticola Rosa villosa Sagina subulata Salix aurita x S. phyllicifolia Salix aurita x S. repens? Salix capraea subsp. sericea Salix lanata Salix lanata x S. lapponum Salix lapponum Salix lapponum x S. repens Salix phylicifolia x S. cinerea Salix phylicifolia x S. repens Salix repens Salix repens x S. herbacea Salix repens x S. myrsinites? Salix repens x S. phylicifolia Saussurea alpina Saxifraga aizoides Saxifraga hypnoides Saxifraga nivalis Saxifraga oppositifolia Schoenus ferrugineus? Schoenus nigricans Sedum roseum Silene acaule? Solidago virgaurea (+ f. cambrica) Taraxacum palustre? Thalictrum alpinum Thymus drucei (form) Trichophorum cespitosum subsp. cespitosum Trichophorum cespitosum subsp. germanicum Trientalis europaea Trifolium repens Trollius europaeus Vaccinium uliginosum Viola riviniana subsp. minor Watsonia 26: 179-195 (2006) 179 Plant Records Records for publication must be submitted to the appropriate Vice-county Recorder (see BSBI Year Book 2005), and not to the Editors. Following publication of the New Atlas of the British & Trish Flora and the Vice-county Census Catalogue, new criteria have been drawn up for the inclusion of records in Plant Records. (See BSBI News 95, January 2004 pp. 10 & 11). These are outlined below: e First records of all taxa (species, subspecies and hybrids) included in the VCCC, designated as native, archaeophyte, neophyte or casual. e First record since 1970 of the taxa above, except in the case of Rubus, Hieracium and Taraxacum. e Records demonstrating the rediscovery of all taxa published as extinct in the VCCC or subsequently. e Newly reported definite extinctions. e Deletions from the VCCC (e.g. through the discovery of errors, the re-determination of specimens etc.) NB — only those errors affecting VCCC entry. e New 10km square records for Rare and Scarce plants, defined as those species in the New Atlas mapped in the British Isles in 100 10km squares or fewer. (See BSBI News 95, January 2004 pp. 36-43). Records for the subdivisions of vice-counties will not be treated separately; they must therefore be records for the vice-county as a whole. However, records will be accepted for the major islands in v.cc. 102-104, 110 and 113. In the following list, records are arranged in the order given in the List of Vascular Plants of the British Isles and its supplements by D.H. Kent (1992), from which the species’ numbers, taxonomy and nomenclature are taken. The Ordnance Survey national grid reference follows the habitat and locality. With the exception of collectors’ initials, herbarium abbreviations are those used in British and Irish Herbaria by D.H. Kent & D.E. Allen (1984). Records are field records if no other source is stated. For all records ‘det.’ or ‘conf.’ appear after the herbarium if the determination was based on material already in an institutional herbarium, otherwise before the herbarium. The following signs are used: * before the vice-county number: to indicate a new vice-county record. + before the species number: to indicate that the plant is an archaeophyte. + before the species number: to indicate that the plant is a neophyte. © before the species number: to indicate that the plant is a casual. The above 3 signs may also used before the vice-county number to indicate the status of the plant in that vice-county. ® before the vice-county number: to indicate that this is an additional hectad for a Rare or Scarce plant. @ at end of entry: established taxon not in Vice-County Census Catalogue. [ ] enclosing a previously published record: to indicate that that record should be deleted or changed. The following list contains records for the year 2005 and earlier. Further records for this period will be published in the next issue of Watsonia. *£2/1.2 SELAGINELLA KRAUSSIANA *4, N. Devon: In garden at Watermouth, SS557480, M. Puttnam, 2005, conf. M. Pool. 4/1.1x3 EQUISETUM XxTRACHYODON (EQUISETUM HYEMALE x E. VARIEGATUM) “*94, Banffs.: Dominant and vigorous on shingle/soil island, 8 m x max 3 m, pure stand by flowing water, less dense on higher part of island, W side of River Avon, NJ158188, A.G. Amphlett, 2005. 4/1.3 EQUISETUM VARIEGATUM *63, S.W. Yorks.: Went Valley, SE649180, S. Yorks CC staff, 1980. 180 PLANT RECORDS 5/1.2 OPHIOGLOSSUM AZORICUM *3, S. Devon: 60+ fertile fronds with infertile fronds at NE end of Greenland Lake, Dawlish Warren, SX985792, N. Mugan & K. Ryland, 2005, det. A.C. Jermy, BM. *94, Banffs.: Growing in damp areas in clearings amongst trees, Spey Bay, NJ357648, IP. Green, 2004. 10/1.1 HYMENOPHYLLUM TUNBRIGENSE 60, W. Lancs.: Grit rocks, Thorn Crag over Wyresdale, SD595572, E.F. Greenwood, 2000, LIV. Originally determined as H. wilsonii but re- determined in 2006. Ist record since 1881. [10/1.2 HYMENOPHYLLUM WILSONIL 37, Worcs.: Error in VCCC - no known records. Delete. ]} 15/2.5a ASPLENIUM TRICHOMANES subsp. TRICHOMANES *60, W. Lancs.: Grit cliffs, Cocklett Scar, Foxdale, Caton-with-Littledale, SD574609, E.F. Greenwood & J.M. Newton, 2004, conf. F.J. Rumsey. Also found in similar habitat at Hawthornthwaite Greave over Wyresdale (SD567517) conf. F.J. Rumsey. 15/2.5c ASPLENIUM TRICHOMANES subsp. PACHYRACHIS *357, Derbys.: Lathkill Dale, SK184658, P. Rushton, 2004. +16/2.1 ONOCLEA SENSIBILIS *94, Banffs.: Large patch in old garden, Glen Wood, Crossbrae, NJ6952, I.P. Green, 2004. 16/5.2 CYSTOPTERIS DICKIEANA *89, E. Perth: Two small colonies 7 m apart on small rock- face forming part of steep bank of burn, Den of Riechip, Dunkeld, NO061488, M.C. Robinson, 2005, conf. F.J. Rumsey, BM. Seen in situ by F.J. Rumsey on 12/8/05. . 16/5.dia CYSTOPTERIS DIAPHANA *3, S. Devon: Flushed banks of old railway cutting, Tavistock, SX475743, R. Hutchins, 2004, det. R. Murphy. Previously recorded as C. fragilis. 17/1.1x2 POLYSTICHUM XBICKNELLII (POLYSTICHUM SETIFERUM x P. ACULEATUM) ES. Derbys.: Three plants at top of steep 2 m bank to river in light woodland and scrub, Lathkill Dale, SK200661, B. Byrne, 2004, det. K. Trewren. 17/3.8x9 DRYOPTERIS XDEWEVERI (DRYOPTERIS CARTHUSIANA X D. DILATATA) *339, Staffs.: Wet, heathy field near Wallbank Grange, west of Cauldon, SK056497, I.C. Trueman & E.V.J. Kohn, 2003. +20/1.nor ABIES NORDMANNIANA *H6, Co. Waterford: Several trees of various sizes self- sown on side of forest ride, Glenabbey, $188199, P.R. Green, 2005. +20/7.mug PINUS MUGO =*©63, S.W. Yorks.: Hugset Wood, SE3006, G. Carr, 1986. +20/7.peu PINUS PEUCE *@©63,S.W. Yorks.: Smiths Wood, Sheffield, SK3286, Sheffield City Ecology Unit, 1988. £21/4.squ JUNIPERUS SQUAMATA *77, Lanarks.: Well established in long abandoned industrial estate, The Gorbals, Glasgow, NS590635, P. Macpherson, 2005, det. M. Gardner, herb. P.M. See BSBI Scottish Newsletter No. 28 page 24. © 28/13.11 RANUNCULUS SCELERATUS 94, Banffs.: One plant growing in a very wet area in landfill site, Nether Dallachy, NJ364641, I.P. Green, 2005. 1st record since 1970. +28/9.hupxvit ANEMONE XHYBRIDA (ANEMONE HUPEHENSIS X A. VITIFOLIA) *©39, Staffs.: A patch in pavement, against a wall, Philip Victor Road, east of Handsworth, SP049904, M.W. Poulton, 2005. *94, Banffs.: Several plants on waste ground, Portsoy, Institute Street, NJ590660, A.G. Amphlett, 2005. Probably cleared house site. +29/1.2 BERBERIS THUNBERGII *57, Derbys.: Roadside hedge, Allestree, SK331395, T. Taylor & B. Rhodes, 2004. Presumably bird-sown origin. +29/1.6 BERBERIS GAGNEPAINII *©57, Derbys.: In pavement under a garden bush on north side of road, Monyash, SK1566, R. Burton, 2005. +29/1.7 BERBERIS JULIANAE *63, S.W. Yorks.: Brecks Plantation, SE6207, G.T.D. Wilmore, 2005. £29/2.1 MAHONIA AQUIFOLIUM *H6, Co. Waterford: Small patch growing out of wall, Waterford, S61661149, D. McGrath, 2005, det. P-.R. Green. Known on this wall by finder for 8—10 years but not named until 2005. £30/1.2 PAPAVER ATLANTICUM *85, Fife: Scrubby sea-bank, by shore track to Hawkcraig, Aberdour, NT196851, G.H. Ballantyne, 2005. Probably escaped from garden 250 yards away. £31/5.1b FUMARIA CAPREOLATA subsp. BABINGTONII *©63, S.W. Yorks.: Building site, Cliffe Road, Bradford, SE167344, B.A. Tregale, 2004. +35/2.1 FICUS CARICA *H6, Co. Waterford: Two large bushes self-sown on face of sea cliff, Tramore, $58060087, P.R. Green, 2005. PLANT RECORDS 181 36/1.1gal URTICA DIOICA subsp. GALEOPSIFOLIA *3, S. Devon: By the Gissage, Honiton, ST1500, L.J. Margetts, 2003, conf. M.F. Godfrey. ©43/1.11 CHENOPODIUM URBICUM _ 57, Derbys.: Weedy vegetable patch, Rodsley, SK205401, G. Hirons, 2003. Ist record since 1970. +43/1.bot CHENOPODIUM BOTRYS *@©3, S. Devon: Garden weed, Trusham, SX853812, R.E.N. Smith, 2001, herb. R.E.N.S. *94, Banffs.: Two plants at landfill site, Nether Dallachy, NJ365643, LP. Green, 2005, det. E.J. Clement. @ ©44/1.1 AMARANTHUS RETROFLEXUS *94, Banffs.: One plant at landfill site, Nether Dallachy, NJ365643, I.P. Green, 2005. ©44/1.cau AMARANTHUS CAUDATUS *94, Banffs.: Several plants at landfill site, Nether Dallachy, NJ365643, I.P. Green, 2005. 46/3.1 HONCKENYA PEPLOIDES 106, E. Ross: Sandy shore, Shandwick, NH859750, B.R. & C.B. Ballinger, 2005. 1st record since 1970. 46/4.2 MINUARTIA VERNA *63, S.W. Yorks.: Levitt Hagg Quarries, SE538009, D.M. Bramley, 1992. 46/5.3 STELLARIA PALLIDA *H6, Co. Waterford: Scattered on dunes, Woodstown, $69880595, P.R. Green, 2005. 46/7.2x3 CERASTIUM XMAUERI (CERASTIUM ARVENSE X C. TOMENTOSUM) *63, S.W. Yorks.: Road verge, Rossington, SK644971, J.G. Hodgson, 1976. 46/10.3 SAGINA SUBULATA — 13, W. Sussex: On and by sandy track and northwards into next tetrad, Flexham Park, TQ007219, F.L. Abraham, 2005, conf. A.G. Knapp. Ist record since c. 1931. *t79, Selkirks.: At 440 m on forest track, above Over Phawhope, Ettrick, NT183072, R.W.M. Corner, 2005, herb. R.W.M.C. Part of Southern Upland Way. Introduced on footwear? 46/10.3x5 SAGINA XMICRANTHA (SAGINA SUBULATA X S. PROCUMBENS) *4, N. Devon: Lundy, SS136456, D. Allen, 2002, det. N. Jardine. 46/17.2 SPERGULARIA MEDIA *©63, S.W. Yorks.: Wardsend Cemetery, SK340902, A. McCarthy, 2000. 46/17.3 SPERGULARIA MARINA *T65, N.W. Yorks.: Edge of A66, Stephen Bank, Scotch Corner, NZ118110, A. Weston, 2000. £46/17.5 SPERGULARIA BOCCONEI 3, S. Devon: Gravel track by the sea, Beesands, SX820407, M. Catt & R.E.N. Smith, 1996, NMW, det. T.C.M. Rich (2005). Ist record since 1936. £46/23.1 VACCARIA HISPANICA *H6, Co. Waterford: Two plants on waste ground, Tramore, $58990267, P.R. Green, 2005, herb. P.R. Green. 46/25.5 DIANTHUS DELTOIDES 89, E. Perth: Several places on stable shingle banks, River Tay at Delvine, NO1238, M.C. Robinson, 2004. +47/3.1 FAGOPYRUM ESCULENTUM *©H6, Co. Waterford: Five plants on waste ground, Tramore, $59390221, P.R. Green, 2005. 47/8.13x15 RUMEX xXSAGORSKII (RUMEX CRISPUS x R. SANGUINEUS) *H6, Co. Waterford: Single plant on waste ground, Snugborough, X07119238, P.R. Green, 2005. 47/8.13x%19 RUMEX XPRATENSIS (RUMEX CRISPUS X R. OBTUSIFOLIUS) 94, Banffs.: Spey Bay, NJ3664, I.P. Green & A.G. Amphlett, 2005. 1st record since 1970. 47/8.14x20 RUMEX XWIRTGENII (RUMEX CONGLOMERATUS x R. PALUSTRIS) *37, Worcs.: Wilden Marsh, SO8274, C.B. Westall & R. Maskew, 2003, det. J.A. Akeroyd. 47/8.21 RUMEX MARITIMUS 80, Roxburghs.: Muddy edge of pond, Barons Folly Loch, Ancrum, NT639264, L.W. Gaskell, 2005, det. R.W.M. Corner, herb. R.W.M.C. Only extant site. Extinct at previously known site. *49/1.mlo PAEONIA MLOKOSEWITSHII *33, E. Gloucs.: Jurassic limestone road cutting on Fosse Way, Broadwell, SP194272, E. Norman, 1998, conf. E.J. Clement. Well established by 27th May 2005 when material collected by M.A.R. & C. Kitchen was confirmed by E.J. Clement. Source: Wild Flower Society Magazine, Autumn 1999 p.26. See also BSBI News 73 p.40. © Apparently 1st record for British Isles. ©53/1.8 MALVA VERTICILLATA *65, N.W. Yorks.: Woodland, Deepgill, East Witton, SE1584, A. Weston, 2000. Growing with other aliens. ©53/2.trm LAVATERA TRIMESTRIS *71, Man.: Levelled landfill sown with cornfield weeds, Ramsey, SC445948, L.S. Garrad, 2000. 182 PLANT RECORDS +53/4.1 ALCEA ROSEA _ 85, Fife: Rocky cliff-face below gardens at Lammerlaws, Burntisland, NT238857, G.H. Ballantyne, 2005. 1st record since 1970. +53/5.1 SIDALCEA MALVIFLORA *@©57, Derbys.: One plant in old railway sidings, Pleasley, SK493639, K. Balkow, 2003. *94, Banffs.: 4 clumps in area of waste ground (probably cleared house site), Portsoy, Institute Street, NJ590660, A.G. Amphlett, 2005. +54/1.fla SARRACENIA FLAVA *3, S. Devon: Colaton Raleigh Common, SY0487, R.E.N. & C.J. Smith, 1999, herb. R.E.N.S. @ Apparently lst record for British Isles. 57/1.9a VIOLA PALUSTRIS subsp. PALUSTRIS *94, Banffs.: Cottown Wood, NJ6852, I.P. Green, 2004. ©60/CUC.mel CUCUMIS MELO *37, Derbys.: Open disturbed ground, Heeley, E of River Sheaf, SK3484, K. Balkow, 2003. Mostly imported soil associated with building new bridge and other developments. +60/CUT.max CUCURBITA MAXIMA “*94, Banffs.: One plant in landfill site, Nether Dallachy, NJ365643, I.P. Green, 2005. t61/1.bal POPULUS BALSAMIFERA *©63, S.W. Yorks.: Abbey Lane Sewage Works, SK538901, J.R. Comley, 1992. t61/1.bsp POPULUS ‘BALSAM SPIRE’ (POPULUS TRICHOCARPA xX P. BALSAMIFERA) *©63, S.W. Yorks.: Edge of borrow pit, SK611945, G. Coles, 2004. 61/2.4x9 SALIX XMOLLISSIMA (SALIX TRIANDRA x S. VIMINALIS) *4, N. Devon: Bow, SS718018, N.F. Stewart, 2004, conf. R.D. Meikle. Confirmed by R.D. Meikle as almost certainly var. hippophaifolia. 61/2.9x16 SALIX XFRIESIANA (SALIX VIMINALIS x S. REPENS) *4, N. Devon: Braunton Burrows, R.D. Meikle, 2003. 61/2.11x14 SALIX xXSTREPIDA (SALIX CINEREA X S. MYRSINIFOLIA) *80, Roxburghs.: Bank above burn, Holm Hill, Newcastleton, NY474879, R.W.M. Corner, 2005, det. R.D. Meikle, herb. R.W.M.C. +61/2.ela SALIX ELAEAGNOS *©63, S.W. Yorks.: Ecclesall Wood, Sheffield, SK3181, Sheffield City Ecology Unit, 1998. +62/5.1 ISATIS TINCTORIA 62, N.E.Yorks.: Dismantled railway line, Robin Hood’s Bay, NZ946052, G. Young, 2003, conf. T.C.G. Rich. Ist record since 1930. 62/12.3 RORIPPA ISLANDICA *4, N. Devon: Upper Tamar Lake, SS2812, Devon Ass. Botany Section & Botanical Cornwall Group, 2003. *80, Roxburghs.: Edge of pond, Barons Folly Loch, Ancrum, NT639264, L.W. Gaskell, 2005, det. T.C.G. Rich, herb. R.W.M.C. Ist confirmed record. +£62/14.5 CARDAMINE PRATENSIS flore pleno *94, Banffs.: Several large patches of plants on east side of railway cutting opposite Tesco’s carpark, Keith & Dufftown Railway, NJ427506, A.G. Amphlett, 2004. Very striking plants in full flower. 62/14.6 CARDAMINE IMPATIENS *©63, S.W. Yorks.: Scabba Wood, SE528017, A. Lister, 1981. +62/14.cor CARDAMINE CORYMBOSA *94, Banffs.: Two plants in garden of 19 Bogmoor Road, Bogmoor, NJ356629, I.P. Green, 2004. Accidental introduction from Christie’s garden centre, Fochabers (v.c.95), where naturalised. |*H12, Co. Wexford: Nursery weed in Coolaught Garden Centre, Coolaught, $871359, P.R. Green, 2005. +£62/20.1 LOBULARIA MARITIMA 85, Fife: Rocky cliff-face below gardens at Lammerlaws, Burntisland, NT238857, G.H. Ballantyne, 2005. Ist record since 1970 and Ist established record (escape from gardens above). 62/23.3x5 COCHLEARIA OFFICINALIS x C. DANICA *4, N. Devon: Base of low cliffs at Buck’s Mill, SS3523, M.F. Godfrey, 2003. £62/31.2 CORONOPUS DIDYMUS 85, Fife: On and by sandy paths, Pettycur Cemetery, Kinghorn, NT265864, G.H. Ballantyne, 2005, herb. G.H.B. Ist record since 1970. [62/34.2x3 BRASSICA XHARMSIANA (BRASSICA NAPUS x B. RAPA) 37, Worcs.: Error in VCCC - no known records. Delete. | ©62/34.2a BRASSICA NAPUS subsp. OLEIFERA *94, Banffs.: One plant growing on rough ground next to 19 Bogmoor Road, Bogmoor, NJ356630, I.P. Green, 2004. +62/34.3b BRASSICA RAPA subsp. CAMPESTRIS *©63, S.W. Yorks.: Near Brinsworth, SK4191, Rotherham Biol. Rec. Centre, 1986. PLANT RECORDS 183 ©62/NES.pan NESLIA PANICULATA 63, S.W. Yorks.: Near Firsby Reservoir, SK4995, Rotherham Biol. Rec. Centre, 1994. Ist record since 1970. ©63/1.0do RESEDA ODORATA *57, Derbys.: Open disturbed ground, Heeley, E of River Sheaf, SK3484, K. Balkow, 2003. Mostly imported soil associated with building new bridge and other developments. +65/6.1 DABOECIA CANTABRICA *94, Banffs.: Good sized plant growing by path in woodland, Winding Walks, Leitch’s Wood, NJ359581, IP. Green, 2005. 65/10.1 ARCTOSTAPHYLOS UVA-URSI 63, S.W. Yorks.: Howden Moors, SK1596, P. Ardron, 1988. 1st record since 1970. 65/13.5 VACCINIUM ULIGINOSUM *57, Derbys.: Two bushes in blanket bog at 513m, near Swains Head, SK131983, D.W. Yalden, 2004, det. S. Edwards. 66/1.1 PYROLA MINOR 37, Worcs.: Woodland, edge of disused railway, Wyre Forest, SO743760, P.L. Reade, 2005, conf. J.J. Day. Last published record 1909. *H12, Co. Wexford: About 10 flowering spikes under birch on dunes, Raven Point, T113240, H. Northridge, 2005. 66/1.2 PYROLA MEDIA @®89, E. Perth: Steep wooded ravine, Gleann Mhairc, N of Blair Atholl, NN 885718, P. Cramb, 2000. @®89, E. Perth: Damp broadleaved woodland/carr adjoining loch, Monk Myre, Blairgowrie, NO2142, L. Farrell, 2004. In flower. 67/1.1a MONOTROPA HYPOPITYS subsp. HYPOPITYS *4, N. Devon: Waste ground at Lower Yelland, SS4832, P. Mullen, 1997. +69/3.1 CYCLAMEN HEDERIFOLIUM *63, S.W. Yorks.: Lawn, Brodsworth Hall, SESO07, C.A. Howes, 1992. 72/2.6 RIBES ALPINUM @®39, Staffs.: A single bush in roadside hedge, West of Knowl Wall, $J851392, I.J. Hopkins, 2005. Native elsewhere in vice-county. £73/1.3 CRASSULA HELMSII *©80, Roxburghs.: Margins of pond, Smailholm village, NT646362, R.W.M. Corner, 2005, herb. R.W.M.C. Introduced. *73/5.4 SEDUM SPECTABILE *85, Fife: One clump, Crook of Devon Moss, NO041002, G.H. Ballantyne, 2005. Has probably persisted for some time. £73/5.7 SEDUM SPURIUM *94, Banffs.: Growing on heaps of soil on old airfield, Bogmoor, NJ358631, I.P. Green & Somerset Rare Plant Group, 2004. +73/5.13 SEDUM SEXANGULARE _ 3, S. Devon: Established on wall, Plympton, SX546551, P. Pullen, 2005. Ist record since 1970. [73/5.19 SEDUM HISPANICUM 37, Worcs.: Error in VCCC - no known records. Delete. ] +73/5.hyb SEDUM HYBRIDUM *94, Banffs.: At least 10 plants growing on old airfield, on east side of road, just south of track to Lower Auchenreath, NJ370637, I.P. Green, 2005. @ +74/1.1xrosxchi ASTILBE XARENDSII (ASTILBE JAPONICA x A. ROSEA x A. CHINENSIS) *H6, Co. Waterford: Two clumps on waste ground, Mapstown, X23449381, P.R. Green, 2005. 74/5.3 SAXIFRAGA NIVALIS 89, E. Perth: Scatter of rocks at 850 m on N facing side of hill top, Meall a’Choire Bhuidhe, Glen Lochsie, NO066709, L. Tucker, 2003, conf. M.C. Robinson. Seen and photographed by MCR on 4/7/05. Ist record in this square (different locality) since 1884 and lst in VC since 1960s. +74/6.1 HEUCHERA SANGUINEA *94, Banffs.: Growing under pine trees on east side of B9104, Bogmoor, NJ3563, I.P. Green, 2004. +74/8.1 TELLIMA GRANDIFLORA *33, E. Gloucs.: Secondary woodland on Jurassic limestone beside A436, SO9215, M.A.R. & C. Kitchen, 2002. 94, Banffs.: Growing in woodland south of cemetery, Bellie Hill, NJ354606, I.P. Green, 2005. Ist record since 1970. £75/3.2 SPIRAEA ALBA *H6, Co. Waterford: Large patch in hedge on north side of N25, Newtown, S42630767, P.R. Green, 2005. £75/3.2x3 SPIRAEA XBILLARDII (SPIRAEA ALBA X S. DOUGLASIT) *H6, Co. Waterford: Small patch in hedge on north side of N25, Newtown, S42630767, P.R. Green, 2005. £75/3.mulxthu SPIRAEA XARGUTA (SPIRAEA MULTIFLORA X S. THUNBERGII) *39, Staffs.: One small bush on side of track, Sheriffhales Common, SJ770124, I.J. Hopkins, 2005. *75/4.1 ARUNCUS DIOICUS *60, W. Lancs.: In dense shade by river in wood, Artle Dale, Caton with Littledale, SD534631, E.F. Greenwood & J.M. Newton, 2005. *75/7.1 KERRIA JAPONICA *63, S.W. Yorks.: Brecks Common, Doncaster, SE6207, G.T.D. Wilmore, 2005. 184 PLANT RECORDS £75/8.6 RUBUS PARVIFLORUS *63, S.W. Yorks.: Sunnybank Urban Nature Reserve, SK3486, Sheffield City Ecology Unit, 1995. 75/8.60 RUBUS LINDLEIANUS *77, Lanarks.: Wood edge, NW of Blackwood, NS7843, B. Simpson & P. Macpherson, 2004. 75/8.77 RUBUS QUESTIERI *10, Wight: Roadside verge outside Firestone Copse, S$Z559915, D.E. Allen, 2005, det. A. Newton, BM, HCMS. 75/8.106 RUBUS INCURVATUS “*12, N. Hants.: Irelands Copse, Bentley, SU7944, D.E. Allen, 2005, HCMS. *H6, Co. Waterford: Rocky outcrop, Ahanaglogh, $36680413, P.R. Green & A.C. Leslie, 2005, det. D.E. Allen. 75/8.117 RUBUS POLYANTHEMUS *77, Lanarks.: NW of Blackwood, NS7843, B. Simpson & P. Macpherson, 2004. 75/8.123 RUBUS SEPTENTRIONALIS *77, Lanarks.: Track side, Thinacre, SW of Larkhall, NS7349, B. Simpson & P. Macpherson, 2003. £75/8.134 RUBUS ARMENIACUS *H6, Co. Waterford: Riverside path, Ballynakill, $6311, P.R. Green & J.C. Wallace, 2005, det. D.E. Allen. 75/8.140 RUBUS ROSSENSIS *10, Wight: One large patch, Parkhurst Forest, SZ479910, D.E. Allen, 2005, HCMS. 75/8.142x321 RUBUS ULMIFOLIUS x R. CAESIUS *H6, Co. Waterford: Hedge on side of lane, Summerville, $62830043, P.R. Green, 2005, det. D.E. Allen. 75/8.144 RUBUS ADSCITUS *H6, Co. Waterford: Roadside hedge, Tallowbridge, X00049433, P.R. Green & A.C. Leslie, 2005, det. D.E. Allen. 75/8.154 RUBUS LANATICAULIS *11, S. Hants.: Carter's Copse, Gosport, SU584001, D.E. Allen et al. 2005, conf. A. Newton, BM. 75/8.163 RUBUS VESTITUS *77, Lanarks.: Near Langlands House, East Kilbride, NS6451, B. Simpson, 2005. 75/8.193 RUBUS MELANODERMIS *H6, Co. Waterford: Along roadside at margin of wood, Ballyheeny Bridge, X12368274, P.R. Green, 2005, det. D.E. Allen. 75/8.225 RUBUS BOTRYEROS *H6, Co. Waterford: Clump on hedge bank of lane, Larbartt’s Bridge, $2515, D.E. Allen, 2001, det. A.L. Newton, BM. 75/8.239 RUBUS INSECTIFOLIUS *63, S.W. Yorks.: Loxley Common, SK3090, Sheffield City Ecology Unit, 1989. 75/8.241 RUBUS LARGIFICUS *12, S. Hants.: Cowleigh Copse, East Green, near Bentley, SU801452, D.E. Allen, 2005, conf. A. Newton, BM, HCMS. 75/8.275 RUBUS ASPERIDENS *10, Wight: Colony above Wootton Creek, Firestone Copse, SZ5590, D.E. Allen, 2005, BM, HCMS. 75/8.305 RUBUS BRITANNICUS *10, Wight: Grammars Copse, Brighstone, SZ415841, D.E. Allen & M.J. Burnhill, 2005, BM, HCMS. 75/8.ava RUBUS AVALONIENSIS *8, S. Wilts.: Margin at crossroads, Barnridge Copse, Bentley Wood, East Grimstead, SU234288, D.E. Allen & D.A. Lewis, 2002, conf. A. Newton, BM, HCMS. *11,S. Hants.: Oakwood fragment, North Copse, Tye, Hayling Island, SU728022, D.E. Allen & A.O. Chater, 1985, BM, conf. R.D. Randall & A. Newton. *12, N. Hants.: Wood margin, Baughurst Common, SU577623, D.E. Allen, 2003, conf. A. Newton, BM, HCMS. 75/8.cae RUBUS CAESARIUS *H6, Co. Waterford: Waste ground, Cappoquin, X09869967, P.R. Green & A.C. Leslie, 2005, det. D.E. Allen. 75/8.int RUBUS INTENSIOR *3, S. Devon: Near Chudleigh, SX8578, R.W. Gould, 1997, det. A. Newton, herb. R.W.G. 75/8.pyd RUBUS PYDARENSIFORMIS *3, S. Devon: Abundant about Kingston and Ringmore, SX64, E.S. Marshall, 1894, BM, det. D.E. Allen. See Watsonia 25: 157—174(2004) for details of this new species. © 75/8.seg RUBUS SEGONTIL *H34, E. Donegal: Common over district of middle Finn valley, west of Ballybofey, C0102 & H0397 etc. D.E. Allen, R. Fitzgerald & P. Hodson, 1992, BM, det. D.E. Allen, conf. A. Newton. Ist record for north half of Ireland. 75/8.vin RUBUS VINDOMENSIS *17, Surrey: Margin of Willey Copse, between Bentley and Farnham, $U812458, D.E. Allen, 2005, BM. See Watsonia 25: 157—174(2004) for details of this new species. © PLANT RECORDS 185 75/9.11 POTENTILLA CRANTZII ®89, E. Perth: Limestone crags at 617 m, Coire a’Ghearraig, Glen Lochsie, NO076698, L. Tucker, 2004. £75/11.2 FRAGARIA MOSCHATA ©63, S.W. Yorks.: Todwick, SK4984, Rotherham Biol. Rec. Centre, 2005. Ist record since 1970. +75/12.1 DUCHESNEA INDICA *©89, E. Perth: Patch 2m x 0:5m outside garden on suburban roadside verge, Perth, NO124227, A. Rutherford, 2005. Photographed by M.C. Robinson on 20/10/2005. 75/13.1 GEUM RIVALE *©71, Man: Garden weed in newish development, Maughold, SC469928, T. Castrin, 2002. 75/13.1x2 GEUM XINTERMEDIUM (GEUM RIVALE xX G. URBANUM) 37, Worcs.: Ancient semi- natural woodland, Twiland Wood and Brook, SO9780, R. Maskew, 2003, conf. C.A. Stace & F. Krahulec. lst record since 1970. Several locations. +£75/13.3 GEUM MACROPHYLLUM *©339, Staffs.: Foot of old wall, outside car park, Rugeley, SK045183, LJ. Hopkins, 2005. Not present in any nearby gardens. +75/18.1 ACAENA NOVAE-ZELANDIAE *85, Fife: Several cast-out plants by woodland path, St Bridget’s Wood, Dalgety Bay, NT170837, J. Cook, 2005, herb. G.H.B. £75/18.2x4 ACAENA ANSERINIFOLIA x A. INERMIS *©37, Worcs.: Sandy grassland, West Midland Safari Park, SO802756, W.A. Thompson, 2003, det. E.J. Clement. *75/18.4 ACAENA INERMIS *©63, S.W. Yorks.: Pavement weed, Thurcroft, SK5088, G.T.D. Wilmore, 2005. [75/19.10a ALCHEMILLA FILICAULIS subsp. FILICAULIS 37, Worcs.: Error in VCCC - no known records. Delete. ] £75/21.7 ROSA “HOLLANDICA’ *94, Banffs.: Two bushes growing on west bank of burn, on west side of the B9104, Bogmoor, NJ357628, I.P. Green, 2004. *75/21.8 ROSA FERRUGINEA *63, S.W. Yorks.: Wall, Thackley, Bradford, SE181383, B.A. Tregale, 2004. 75/21.13x17 ROSA XGLAUCOIDES (ROSA CAESIA subsp. VOSAGIACA x R. MOLLIS) 77) Lanarks.: Roadside verge, by hedge, Stonehouse to Udston road, NS751451, P. Macpherson, 2004, det. R. Maskew, herb. P.M. £75/22.1 PRUNUS PERSICA *@©63, S.W. Yorks.: Near canal, Sheffield, SK398907, K. Balkow, 2005. £75/22.2 PRUNUS DULCIS *©63, S.W. Yorks.: Wadsley & Loxley Common, SK3190, Sheffield City Ecology Unit, 2002. 75/22.4x5 PRUNUS XFRUTICANS (PRUNUS SPINOSA X P. DOMESTICA) *94, Banffs.: One bush growing amongst trees on bank on east side of the B9104, Bellie Hill, NJ352610, I.P. Green, 2005. £75/22.ser PRUNUS SERRULATA *@©63, S.W. Yorks.: Burngreave, SK3689, Sheffield Wildlife Trust, 2003. *75/27 miexatr MALUS XPURPUREA (MALUS NIEDZWETZKYANA X M. ATROSANGUINEA) *© 37, Worcs.: Single bush, considered bird sown, Harvington Island, River Avon, SP064477, A.O. Chater & R. Maskew, 2004. 75/28.1x9 SORBUS XTHURINGIACA (SORBUS AUCUPARIA x S. ARIA) *37, Worcs.: Scrub woodland, Malvern Hills, SO773461, P.G. Garner, 2004, det. P.J.M. Nethercott. *©63, S.W. Yorks.: Rokeby Road, Sheffield, SK356929, Sheffield Wildlife Trust, 2004. *75/28.7 SORBUS INTERMEDIA *H6, Co. Waterford: Self-sown on top of wall, Waterford, $61531126, P.R. Green, 2005. *75/30.1 AMELANCHIER LAMARCKII *©63, S.W. Yorks.: Disused railway embankment, Shipley, SE157381, B.A. Tregale, 2004. *£75/32.15 COTONEASTER CONGESTUS *77, Lanarks.: Well established in long abandoned industrial estate, The Gorbals, Glasgow, NS5963, P. Macpherson, 2005, det. J. Fryer, herb. P.M. *75/32.18 COTONEASTER CONSPICUUS *60, W. Lancs.: Self seeded on wall top, car park, Preston, SD533289, M. Wilcox, 2005. *75/32.21 COTONEASTER HORIZONTALIS 94, Banffs.: Cemetery, Fordyce, NJ555638, I.P. Green, 2005. 1st record since 1970. £75/32.24 COTONEASTER ADPRESSUS *©63, S.W. Yorks.: Parkin Wood, Chapeltown, SK3597, K. Dulieu, 2002. 186 PLANT RECORDS +75/32.26 COTONEASTER DIVARICATUS *85, Fife: Several bushes in hedge forming part of boundary, Dunnikier Golf Course, Kirkcaldy, NT273947, G.H. Ballantyne, 2005. Originally planted, probably now self-sowing. *£75/32.37 COTONEASTER DIELSIANUS *©37, Derbys.: Disused railway bed, Long Eaton, SK491344, R.D. Martin, 2005, det. A. Willmot. *75/32.40 COTONEASTER STERNIANUS *13, W. Sussex: Old railway bank, Mid Lavant, SU858084, R.C. Stern, 2005. +75/32.his COTONEASTER HSINGSHANGENSIS *63, S.W. Yorks.: Field, Reynolds Wood, Shipley, SE144361, B.A. Tregale, 2003. +75/32.r0s COTONEASTER ROSEUS *29, Cambs: Several birdsown shrubs at base of railings by old car park for railway station, Newmarket, TL644626, A.C. Leslie & P.R. Green, 2005, det. J. Fryer, CGE. @ Apparently 1st record for the British Isles. *75/33.2 PYRACANTHA ROGERSIANA *©63, S.W. Yorks.: Denholme Close, Sheffield, SK3588, Sheffield Wildlife Trust, 2004. *77/1.1 ROBINIA PSEUDOACACIA *94, Banffs.: In hedge on south side of road, Crossbrae, NJ6952, I.P. Green, 2004. *77/14.ben VICIA BENGHALENSIS ©63, S.W. Yorks.: Old Moor wetland site, Wombwell, SE4202, G.T.D. Wilmore, 2005. 1st record since 1970. £77/14.fab VICIA FABA *71, Man: Amenity site, Ramsey, SC444948, L.S. Garrad, 1999. £77/15.11 LATHYRUS HIRSUTUS ©63,S.W. Yorks.: Cannon Hall, Barnsley, SE2708, Barnsley MBC records, 1991. Ist record since 1970. £77/15.7 LATHYRUS GRANDIFLORUS *H6, Co. Waterford: Patch on edge of scrub opposite Murphy’s Bar, NCR, Clohernagh, S63750244, P.R. Green, 2005, herb. P.R. Green. ©77/15.ann LATHYRUS ANNUUS *71, Man: Andreas tip, SC433987, M. Devereau, 1980. Probably from pig farm waste. 77/18.4 MEDICAGO POLYMORPHA ©37, Derbys.: Waste ground, Britannia Mill, Derby, SK342369, S. Jackson, 2004, DBY. Growing on hardcore deposited during building work from down south. Ist record since 1970. *77/19.19a TRIFOLIUM INCARNATUM subsp. INCARNATUM *H6, Co. Waterford: Four plants on waste ground, Tramore, $59150245, P.R. Green, 2005, herb. P.R. Green. 77/19.19b TRIFOLIUM INCARNATUM subsp. MOLINERII 3, S. Devon: Locally abundant on cliff edge, Bolt Tail, SX674389, P. Reay & W. Rees, 2005, herb. R.E.N.S. Ist record since 1900. *77/21.1x2 LUPINUS XREGALIS (LUPINUS ARBOREUS x L. POLYPHYLLUS) *94, Banffs.: Growing in clearing in trees next to rubbish dump, Nether Dallachy, NJ363645, I.P. Green, 2004. £77/22.2 LABURNUM ALPINUM 94, Banffs.: In hedge on S. side of road, Crossbrae, NJ6952, I.P. Green, 2004. Ist record since 1970. é ©77/DOR.hir DORYCNIUM HIRSUTUM *39, Staffs.: One plant in pavement against wall by 83 Jayshaw Avenue, Great Barr, SP047938, M.W. Poulton, 2005, conf. E.J. Clement. @ £78/2.mac ELAEAGNUS MACROPHYLLA *3, S. Devon: By coast path at St Mary’s Bay, SX9355, R.E.N. & C.J. Smith, 2005, herb. R.E.N.S. +78/2.macxpun ELAEAGNUS XSUBMACROPHYLLA (ELAEAGNUS MACROPHYLLA xX E. PUNGENS) *©63, S.W. Yorks.: Thorpe Marsh Nature Reserve, SE5909, B.K. Byrne, 2002. 84/1.1x8 EPILOBIUM XNOVAE-CIVITATIS (EPILOBIUM HIRSUTUM x E. CILIATUM) *60, W. Lancs.: Car park, Preston, SD536288, M. Wilcox, 2005. 84/1.5 EPILOBIUM TETRAGONUM *60, W. Lancs.: Waste ground, Preston, $D533292, M. Wilcox, 2005. Ist confirmed v.c record. Specimen provided by P. Jepson to be deposited at LIV. +84/4.5 OENOTHERA STRICTA 33, E. Gloucs.: Waste ground under development, north side of A4019, near Tewkesbury Bridge, Cheltenham, $0934238, M.A.R. & C. Kitchen, 2005. 1st record since 1970. 91/2.12 EUPHORBIA PORTLANDICA 13, W. Sussex: Coastal shingle, patch 2m square on top of sea wall facing Hayling Island, S. Thorney Island, SU761014, Sussex Botanical Recording Society, 2005, conf. A.G. Knapp. Ist record since 1963. +91/2.6 EUPHORBIA PLATYPHYLLOS *@©57, Derbys.: In taller growth on disturbed ground, M1 Junction 25 south-west side, SK471354, G. Hirons, 2004. £91/2.9 EUPHORBIA LATHYRIS *94, Banffs.: One plant in landfill site, Nether Dallachy, NJ365643, I.P. Green, 2005. PLANT RECORDS 187 +91/2.0bl EUPHORBIA OBLONGATA *@©63, S.W. Yorks.: Rough ground, St. Luke’s, Bradford, SE158317, B.A. Tregale, 2005. £93/2.3 PARTHENOCISSUS TRICUSPIDATA *63, S.W. Yorks.: Shirland Lane, Sheffield, SK3888, Sorby Natural History Society, 1988. £94/1.2 LINUM USITATISSIMUM 94, Banffs.: Several plants in landfill site, Nether Dallachy, NJ365643, I.P. Green, 2005. Ist record since 1970. £102/1.3 OXALIS CORNICULATA *94, Banffs.: Growing as a weed in the garden of a house in the village of Bogmoor, NJ3563, I.P. Green, 2004. +103/1.13xpla GERANIUM XMAGNIFICUM (GERANIUM IBERICUM x G. PLATYPETALUM) *H6, Co. Waterford: Clump on road verge, Kilcana, $38881047, P.R. Green, 2005, herb. P.R. Green. £103/1.2 GERANIUM VERSICOLOR *65, N.W. Yorks.: By road south of Bland’s Gill, north of Sedburgh, SD634949, C.E. Wild, 2005. +103/2.2 ERODIUM MOSCHATUM ©39, Staffs.: Grassland, tens of plants between rear boundary fence of Sports Stadium and lake, Perry Barr Park, SP066926, J.E. Hawksford, 2005, STOKE. First record since 1906. Possibly accidentally introduced with sand used for field events on the other side of the fence. *©57, Derbys.: Open disturbed ground, Heeley, E of River Sheaf, SK3484, K. Balkow, 2003. Mostly imported soil associated with building new bridge and other developments. +103/LIM.dou LIMNANTHES DOUGLASII *94, Banffs.: Spreading along 5 metres of road verge, opposite houses, NE of Moss of Bourach, NJ255405, A.G. Amphlett, 2005. ©104/1.maj TROPAEOLUM MAJUS *94, Banffs.: Growing in trees opposite houses in Bogmoor, E side of the B9104, NJ3563, I.P. Green, 2004. +107/1.ran HYDROCOTYLE RANUNCULOIDES *60, W. Lancs.: Marl pit, Bispham, SD326408, M. Sutcliffe, 2005. *107/21.1 FOENICULUM VULGARE *94, Banffs.: Two plants growing on edge of rubbish tip, just where it joins the old airfield. Moor of Dallachy, NJ369638, I.P. Green, 2005. 107/31.1 CICUTA VIROSA ®13, W. Sussex: Bank of River Rother, well out of water, near South Grounds Farm, Stopham, TQ024175, F.L. Abraham, 2005. Also present at two other places - TQ028175 & TQ028176. 108/3.2x4 CENTAURIUM ERYTHRAEA X C. PULCHELLUM *37, Worcs.: Forestry ride, Tiddesley Wood, SO928457, A.W. Reid, 2002, conf. F. Ubsdell. 108/5.4a GENTIANELLA AMARELLA subsp. AMARELLA *94, Banffs.: 69 plants in two areas of dry calcareous grassland, grazed by rabbits with much bare ground, at a height of 450 m in Glen Builg, NJ181064 and NJ182062, A.G. Amphlett, I.P. Green, G. Roberts & R. Finch, 2005. Plants in bud, no flowers fully open. +110/6.1 PHYSALIS ALKEKENGI *339, Staffs.: More than 30 plants in waste ground resulting from housing demolition, Foxoak Street, Dudley Wood, SO944861, M.W. Poulton & J.E. Hawksford, 2005. £110/6.per PHYSALIS PERUVIANA * 3, S. Devon: On compost pile, Poolmill Cross, SX807820, N.F. Stewart, 2003. +110/9.1 DATURA STRAMONIUM *94, Banffs.: Two plants in landfill site, Nether Dallachy, NJ365643, LP. Green, 2005. +110/NIC.alaxfor NICOTIANA XSANDERAE (NICOTIANA ALATA xX N. FORGETIANA ) *©39, Staffs.: 1 plant in waste ground resulting from housing demolition, St. Anne’s Road, Cradley Heath, SO942859, M.W. Poulton & J.E. Hawksford, 2005. *94, Banffs.: Several plants in landfill site, Nether Dallachy, NJ365643, I.P. Green, 2005. ©110/PET.axixint PETUNIA XHYBRIDA (PETUNIA AXILLARIS x P. INTEGRIFOLIA) “O)// Derbys.: Open disturbed ground, Heeley, E of River Sheaf, SK3484, K. Balkow, 2003. Mostly imported soil associated with building new bridge and other developments. 111/3.2x4 CALYSTEGIA XLUCANA (CALYSTEGIA SEPIUM xX C. SILVATICA) 3, S. Devon: Northcote Lane, Honiton, ST1500, L.J. Margetts, 2005, conf. R.K. Brummitt, K. 1st record since ROTO: 111/3.3x4 CALYSTEGIA XHOWITTIORUM (CALYSTEGIA PULCHRA X C. SYLVATICA *63, S.W. Yorks.: Marsden, Huddersfield, SEO519, M.J. Lucas, 2002. 188 PLANT RECORDS +111/3.4dis CALYSTEGIA SILVATICA subsp. DISJUNCTA *3, S. Devon: By River Gissage, Honiton, ST1500, L.J. Margetts, 2000, det. R.K. Brummitt. © +114/1.pau POLEMONIUM PAUCIFLORUM *©39, Staffs.: Half a dozen plants scattered in waste ground resulting from housing demolition, Newtown Lane, Dudley Wood, SO946862, M.W. Poulton & J.E. Hawksford, 2005, STOKE. © *115/1.1 PHACELIA TANACETIFOLIA *4, N. Devon: Braunton, SS43, M. Breeds, 2005. 116/2.1 ECHIUM VULGARE +94, Banffs.: Two plants growing on the edge of grass as it meets the shingle, just east of Spey Bay Hotel, NJ355652, I.P. Green, 2005. Ist record since 1970. 7116/2.2 ECHIUM PLANTAGINEUM *@©57, Derbys.: Open disturbed ground, Heeley, E of River Sheaf, SK3484, K. Balkow, 2003. Mostly imported soil associated with building new bridge and other developments +116/2.4 ECHIUM PININANA *©H12, Co. Wexford: Single plant self-sown at base of wall on roadside, Millroad, S977044, P.R. Green, 2005. *116/4.6 SYMPHYTUM ORIENTALE *@©80, Roxburghs.: Wooded side of picnic area by A7 at Teviothead, NT402041, R.W.M. Corner, 2005, herb. R.W.M.C. Presumed to have been planted. +116/5.1 BRUNNERA MACROPHYLLA *94, Banffs.: 1 plant growing on rough ground on north side of track on the east edge of Nether Dallachy, NJ3663, I.P. Green, 2004. ©116/6.3 ANCHUSA AZUREA *13, W. Sussex: one plant on dumped soil at 185m altitude, Summer Down, TQ265108, A. Spiers, 2005. 116/15.3 MYOSOTIS STOLONIFERA ®79, Selkirks.: A small population in flower in flush at — 265m, N of Shorthope Sike, Deloraine Burn, Ettrick, NT338199, R.W.M. Corner, 2005, herb. R.W.M.C. ©116/CER.maj CERINTHE MAJOR *57, Derbys.: Waste ground bank, Station Road, Sandiacre,. SK480363, R.D. Martin, 2005, det. A. Willmot. @ +118/l.ann STACHYS ANNUA 17, Surrey: Disused golf course, Chessington, TQ196636, C.L. Coleman & J. O’Reilly, 2004, conf. E.J. Clement, herb. C.O.R. & J.O.R. Ist record since 1970. Probably Ist national record for c. 20 years. +118/2.pse BALLOTA PSEUDODICTAMUS *3, S. Devon: On bank near church at Cornworthy, SX8255, L.J. Margetts & L.M. Spalton, 1996. © *118/7.1 PHLOMIS RUSSELIANA *33, E. Gloucs.: Patch 2m x 3m in small, overgrown limestone quarry, Arlington, Bibury, SP108070, M.A.R. & C. Kitchen, 2005, conf. E.J. Clement. Away from buildings. Theatened by dumping. £118/7.2 PHLOMIS FRUTICOSA *34, W. Gloucs.: Disused Jurassic limestone quarry, Breakheart Hill, North Nibley, ST756967, J.R. Doe, 2004. £118/10.1 SCUTELLARIA ALTISSIMA *257, Derbys.: Old railway land, Ilkeston, SK462425, B. Gough, 2005, det. B.S. Wurzell. Known here since 2002. 118/15.1x2 PRUNELLA XINTERMEDIA (PRUNELLA VULGARIS x P. LACINIATA) 37, Worcs.: Limestone grassland, Penny Hill, SO750612, R. Maskew, 1991, conf. W.A. Thompson. Ist record since 1970. *118/16.1 MELISSA OFFICINALIS *65, N.W. Yorks.: Woodland, Throstle Gill, Dalton, NZ1108, A. Weston, 2000. Also found shortly after beside path in an open calcareous site in Leyburn Shawl Wood (SE099) by D.J. Millward. *94, Banffs.: One plant growing in landfill site, Nether Dallachy, NJ363641, I.P. Green, 2005. 118/23.5 MENTHA PULEGIUM ®13, W. Sussex: Bare mud at edge of small lake (several plants around GR on N side, abundant on S side of lake), East of Bolney, TQ269227, A.G. Knapp, 2005. Appears native but may be introduction. *118/24.1 ROSMARINUS OFFICINALIS *H6, Co. Waterford: Self-sown on side and at base of wall, Ballynakill, S638109, D. McGrath, 2005. *118/25.1 SALVIA SCLAREA *©39, Staffs.: Seedlings in gaps between flagstones, Common Road, Wombourne, SO0872924, C.B. Westall, 2005. *122/1.1 BUDDLEJA ALTERNIFOLIA *©63, S.W. Yorks.: Burngreave, Sheffield, SK3689, Sheffield Wildlife Trust, 2003. ©122/1.3 BUDDLEJA GLOBOSA *13, W. Sussex: Ifield Station, TQ251366, G. Bursnall, 2005. *124/1.10 VERBASCUM SPECIOSUM *17, Surrey: Roadside verge, Cobham, TQ118617, C.L. Coleman & J. O’Reilly, 2004, conf. V. Johnstone, herb. C.O.R. & J.O.R. PLANT RECORDS 189 +124/4.1 MIMULUS MOSCHATUS *H6, Co. Waterford: Plentiful in marshy area, Strancally Quay, X08539038, P.R. Green & M.J. Stribley, 2005. [124/4.3xcup MIMULUS XMACULOSUS (MIMULUS LUTEUS x M. CUPREUS) 37, Worcs.: Error in VCCC - no known records. Delete. ] 124/5.1 LIMOSELLA AQUATICA ®39Staffs.: 1000s of plants in mud surround of reservoir drained for repair, Knypersley Reservoir, SJ894555, E. Oxenham, 2005. £124/7.1 ANTIRRHINUM MAJUS *94, Banffs.: Growing in bare areas amongst moss on old airfield, on south bank of track to Lower Auchenreath, NJ370638, I.P. Green, 2005. £124/14.2 DIGITALIS LUTEA “*85, Fife: About ten plants in scrubby waste ground, shore, west of old harbour, Buckhaven, NT352977, G.H. Ballantyne, 2005. Possibly introduced with a seed- mix. 124/16.11x12 VERONICA ™XLACKSCHEWITZII (VERONICA ANAGALLIS-AQUATICA x V. CATENATA) *H6, Co. Waterford: Single plant on edge of pool, Portnaboe, $43692070, P.R. Green, 2005. £124/16.25 VERONICA LONGIFOLIA *94, Banffs.: Growing on side of track near Nether Dallachy, NJ362638, I.P. Green, 2004. 124/16.25x26 VERONICA LONGIFOLIA x V. SPICATA *©63, S.W. Yorks.: Cowper Ave., Sheffield, SK337932, Sheffield City Ecology Unit, 1991. £124/16.26 VERONICA SPICATA *©63, S.W. Yorks.: Brightside goit side, Sheffield, SK383891, O.L. Gilbert, 1988. +124/17.1xell HEBE XLEWISII (HEBE SALICIFOLIA x H. ELLIPTICA) *H6, Co. Waterford: Self- sown on side of wall, Dunmore East, $689000, P.R. Green, 2005. Parent plant in nearby garden. [124/20.3 EUPHRASIA ANGLICA 37, Worcs.: Date-class error in VCCC; last recorded in 1961.] ®39, Staffs.: Heathy, hilly pasture between Lud’s Church and Back Dane, SJ979661, IJ. Hopkins, 2005. A nationally endangered species. 124/20.8 EUPHRASIA PSEUDOKERNERI *63, S.W. Yorks.: Calcareous grassland, Kiveton Park, SK5082, W.A. Ely, 2004, conf. J.R. Comley. £129/1.5 CAMPANULA MEDIUM *©H6, Co. Waterford: Single plant on waste ground, Ballynacourtey, X27378963, P.R. Green, 2005. *£129/1.8 CAMPANULA PYRAMIDALIS *71, Man: Relic in derelict garden, Peel, SC247839, L.S. Garrad, 2000. +£129/1.9 CAMPANULA PORTENSCHLAGIANA *63, S.W. Yorks.: Wall, Lynton Road, Sheffield, SK3385, K. Balkow, 2005. +129/7.2 LOBELIA ERINUS *94, Banffs.: One plant growing by track in pinewood, Spey Bay, NJ358646, I.P. Green, 2004. £131/5.1 LEYCESTERIA FORMOSA *79, Selkirks.: A strong colony on rubble dumped to stabilise river bank, River Ettrick upstream from Selkirk Bridge, NT460282, D. Methven, 2005, det. D. McKean, E. £131/6.1 LONICERA PILEATA *3, S. Devon: Self-seeded by main carpark, Honiton, ST1600, L.J. Margetts, 2005, herb. R.E.N.S. *60, W. Lancs.: Waste ground, Preston, SD533292, M. Wilcox, 2005. *94, Banffs.: One small bush growing on west side of path by entrance to Fife Keith Wood, NJ416507, I.P. Green, 2005. Might have been planted. *131/6.2 LONICERA NITIDA *94, Banffs.: One bush on north side of track, just south of the A98, Red Slack, NJ363588, IP. Green, 2005. £131/6.6 LONICERA JAPONICA *4, N. Devon: Garden weed, Torrington, SS5018, W.H. Tucker, 2005. *60, W. Lancs.: Bank overlooking brackish pool, Fleetwood Nature Park, SD335465, E.F. Greenwood, 2005. +131/WEI.flo WEIGELA FLORIDA *©63,S.W. Yorks.: Nottingham Street, Sheffield, SK3588, R. Butterfield, 2003. £133/3.1 CENTRANTHUS RUBER *94, Banffs.: Two plants growing on old airfield, Moor of Dallachy, NJ369638, I.P. Green, 2005. £134/1.2 DIPSACUS SATIVUS *71, Man: Waste ground, Ramsey, SC445946, L.S. Garrad & P. Davey, 2000. £134/2.1 CEPHALARIA GIGANTEA 3, S. Devon: Three plants established on roadside at Arnold’s Point, Plymouth, SX503553, P. Pullen, 2005. *94, Banffs.: Growing on heaps of soil on the old airfield. Bogmoor, NJ358631, I.P. Green & Somerset Rare Plant Group, 2004, det. P.R. Green. 190 PLANT RECORDS +134/3.mac KNAUTIA MACEDONICA *71, Man: Gravelled access, Ramsey, SC446954, L.S. Garrad, 2000. @ Apparently Ist record for the British Isles. +135/1.3 ECHINOPS BANNATICUS *33, E. Gloucs.: Road verge, Ford, SP086292, J.P. Martin, 2005. 135/11.3 CENTAUREA CYANUS +94, Banffs.: Several plants in landfill site, Nether Dallachy, NJ365643, I.P. Green, 2005. 1st record since 1970. ©135/11.6 CENTAUREA SOLSTITIALIS *37, Derbys.: Disturbed ground on bank of newly- excavated pond, Barlow, SK347753, K. Balkow, 2003, DBY. Farmer had deliberately spread aviary sweepings. +135/12.1 CICHORIUM INTYBUS 93, N. Aberdeen: Old grassland in small field, Balhalgardy, Inverurie, NJ757242, A. Watson, 2002, herb. D. Welch. Ist record since 1930. 135/16.3 LEONTODON SAXATILIS *£94, Banffs.: At least 40 plants growing on old airfield, on east side of road, just north of track to Lower Auchenreath, Moor of Dallachy, NJ369638, I.P. Green, 2005. 135/24.1 MYCELIS MURALIS *+94, Banffs.: One plant on side of path in pine wood opposite Cunninghaugh, NJ356646, I.P. Green, 2004. 135/25.22 TARAXACUM RUBICUNDUM 33, E. Gloucs.: Bredon Hill, Kemerton, SO0959402, A.W. Reid, 2005. Ist record since 1970. 135/25.31 TARAXACUM ANGLICUM *37, Worcs.: Unimproved mesic grassland, Chaceley Meadows SSSI, SO855305, M.V. Marsden, 1987, det. C.C. Haworth. 135/25.66 TARAXACUM HESPERIUM *37, Worcs.: Disturbed ground, Droitwich Railway Station, SO893633, J.J. Day, 2005, det. A.J. Richards & A.W. Reid, herb. A.W. Reid. 135/25.68 TARAXACUM INANE *37, Worcs.: Damp grassland, Kingswood Common, SO745603, A.W. Reid & R. Maskew, 2003, det. A.J. Richards, herb. A.W. Reid. 135/25.73 TARAXACUM OELLGAARDII *37, Worcs.: Unimproved mesic grassland, Chaceley Meadows SSSI, SO856305, A.W. Reid & K. Barnett, 2005, det. A.J. Richards & A.W. Reid, herb. A.W. Reid. 135/25.78 TARAXACUM PORTERI *37, Worcs.: Laneside bank, Battenton Green, SO828659, J.J. Day, 2004, det. A.J. Richards, herb. A.W. Reid. 135/25.81 TARAXACUM TAMESENSE *37, Worcs.: Unimproved mesic grassland, Chaceley Meadows SSSI, S0855305, M.V. Marsden, 1988, det. C.C. Haworth. 135/25.91 TARAXACUM KERNIANUM *37, Worcs.: Grass verge, Lower Shepley Lane, Burcot, S0980724, J.J. Day, 2003, det. A.J. Richards, herb. A.W. Reid. £135/25.96 TARAXACUM PRIONUM *37, Worcs.: Laneside bank, Cur Lane, Bentley, SP010699, J.J. Day, 2003, det. A.J. Richards, herb. A.W. Reid. 135/25.99 TARAXACUM QUADRANS _*37, Worcs.: Laneside bank, Elmley Lovett to Podmoor, S0865711, J.J. Day, 2003, det. A.J. Richards, herb. A.W. Reid. £135/25.101 TARAXACUM SUBHAMATUM *63, S.W. Yorks.: Ashbourne Drive, Bradford, SE169352, B.A. Tregale, 2005, conf. A.J. Richards. *135/25.112 TARAXACUM ANGULARE *37, Worcs.: Laneside bank, Broomhall, Kemspey, SO866508, A.W. Reid, 2005, det. A.J. Richards & A.W. Reid, herb. A.W. Reid. *£135/25.127 TARAXACUM DENSILOBUM *33, E. Gloucs.: Road verge, Grafton, Beckford, S0988372, A.W. Reid, 2005, conf. A.J. Richards. +135/25.134 TARAXACUM EXSERTIFORME *37, Worcs.: Grass verge, bridleway, Cames Coomb, Bredon Hill, SO0970415, A.W. Reid, 2005, det. A.J. Richards & A.W. Reid, herb. A.W. Reid. £135/25.135 TARAXACUM EXSERTUM *37, Worcs.: Grass verge, Hulls Lane, Oddingley, S0904599, J.J. Day, 2003, det. A.J. Richards, herb. A.W. Reid. 135/25.139 TARAXACUM HEXHAMENSE *37, Worcs.: Grass verge, Kingsford Lane, Wolverley, S0823799, J.J. Day, 2005, det. A.J. Richards, herb. A.W. Reid. £135/25.141 TARAXACUM HUELPHERSIANUM _*37, Worcs.: Grass verge, Beacon Lane, Lickey, S0979752, A.W. Reid, 2005, det. A.J. Richards & A.W. Reid, herb. A.W. Reid. £135/25.146 TARAXACUM LACERIFOLIUM *63, S.W. Yorks.: Footpath, Shipley, SE157381, B.A. Tregale, 2005, conf. A.J. Richards. 135/25.148 TARAXACUM LACINIOSUM *3, S. Devon: Grass verge near Strete Raleigh, SY0695, L.J. Margetts, 2003, det. A.J. Richards, herb. L.J.M. PLANT RECORDS 191 £135/25.155 TARAXACUM LEUCOPODUM *33, E. Gloucs.: Road verge, Pebworth, SP127467, A.W. Reid, 2005, conf. A.J. Richards. £135/25.162 TARAXACUM MACROLOBUM *37, Worcs.: Grass verge, Lane by Eldersfield Church, SO799311, A.W. Reid, 2005, det. A.J. Richards, herb. A.W. Reid. 135/25.163, TARAXACUM MACULATUM *3, S. Devon: Garden weed, Honiton, ST1500, L.J. Margetts, 2003, conf. A.J. Richards, herb. L.J.M. +135/25.175 TARAXACUM OCHROCHLORUM *63, S.W. Yorks.: Ashbourne Drive, Bradford, SE169352, B.A. Tregale, 2005, conf. A.J. Richards. £135/25.187 TARAXACUM PORRIGENS *37, Worcs.: Grass verge, Broadwas Church, SO0754550, A.W. Reid, 2005, det. A.J. Richards & A.W. Reid, herb. A.W. Reid. +135/25.208 TARAXACUM SUBLONGISQUAMEUM = Sim eWOLES-.8 Werb) top, Parka Gate; $0935715, J.J. Day, 2005, det. A.J. Richards, herb. A.W. Reid. £135/25.211 TARAXACUM SUBXANTHOSTIGMA *©37, Worcs.: Grass verge, Market Street, Bromsgrove, SO959709, J.J. Day, 2004, det. A.J. Richards, herb. A.W. Reid. 135/25.217 TARAXACUM UNDULATUM = *3, S. Devon: Roadside near Strete Raleigh, SY0695, L.J. Margetts, 2003, det. A.J. Richards, herb. L.J.M. +135/27.3a PILOSELLA FLAGELLARIS subsp. FLAGELLARIS *63, S.W. Yorks.: Lindrick Dale, SK540821, J.G. Hodgson, 1976. £135/28.138 HIERACIUM GRANDIDENS *63, S.W. Yorks.: Chellow Dene, Bradford, SE122347, B.A. Tregale, 2004, conf. V. Jones. £135/28.140 HIERACIUM EXOTERICUM agg. *63, S.W. Yorks.: Broom Vale, Sheffield, SK270901, Sheffield City Ecology Unit, 1986. 135/28.arg HIERACIUM ARGILLACEUM *77, Lanarks.: Track side, Uddingston, NS7060, P. Macpherson, 2004, det. D.J. Mcosh, herb. P.M. @ *135/28.med HIERACIUM MEDIUM *3, S. Devon: On wall near old railway, Tavistock, SX4774, C. West, 1952, CGE, det. P.D. Sell. @ Apparently 1st published record for the British Isles. +¢135/28.mon HIERACIUM MONSTROSUM = *3, S. Devon: On wall near old railway, Tavistock, SX4774, C. West, 1952, CGE, det. P.D. Sell. @ Apparently 1st published record for the British Isles. £135/29.1 GAZANIA RIGENS *H6, Co. Waterford: Single plant on edge of dunes, Dungarvan, X277937, P.R. Green, 2005. 135/30.2 FILAGO LUTESCENS @®13, W. Sussex: Several plants in sandy arable field, South Ambersham, SU913205, F.L. Abraham, 2005. +135/33.6 GNAPHALIUM LUTEOALBUM *@©37, Worcs.: Bare ground between path and lawn, Homer Hill, Cradley, SO940849, M.W. Poulton, 2003, conf. F. Rose. Still present in 2004. £135/35.1 INULA HELENIUM *94, Banffs.: Growing on the east bank of the railway line, Keith, NJ428507, LP. Green, 2005. First seen in 2004. +135/38.1 TELEKIA SPECIOSA *94, Banffs.: Large clump in flushed grassland above footpath, Drumin Tower, NJ1830, A.G. Amphlett, 2005, conf. I.P. Green. First seen in 2004 but not identified at the time. +135/40.3 SOLIDAGO CANADENSIS *80, Roxburghs.: S bank of River Teviot, at Ashybank W of Denholm, NT551180, R.W.M. Corner, 1997, herb. R.W.M.C. Scabrid plants with few disc florets per flower. 135/41.pun ASTER PUNICEUS *£94, Banffs.: Patch growing on the waste ground between 19 & 15 Bogmoor Road, Bogmoor, NJ357629, I.P. Green, 2005. © £135/44.1 CONYZA CANADENSIS *87, W. Perth: Landscaped colliery waste, old pit bing, Blairhall, NS999885, G.H. Ballantyne, 2005, herb. G.H.B. Provisionally identified near here in 1979 so may have been present since then. Although in Fife, the locality is in v.c. 87 +£135/44.2 CONYZA SUMATRENSIS *H6, Co. Waterford: Single plant on waste ground, Waterford, S602111, P.R. Green, 2005, herb. P.R. Green. £135/44.bil CONYZA BILBAOANA *94, Banffs.: One plant in flower bed in the garden of 19 Bogmoor Road, Bogmoor, NJ356629, I.P. Green, 2005, det. E.J. Clement. Unknown origin. ©135/44.dav CONYZA DAVEAUANA = *13, W. Sussex: Two plants at base of wall in private car park, Hove, TQ290059, A. Spiers, 2002, det. P.D. Sell, CGE. @ Apparently 1st record for the British Isles. 192 PLANT RECORDS £135/48.coc TANACETUM COCCINEUM *71, Man: Garden dump on development site, Ballagarey, SC341789, L.S. Garrad, 1998. [£135/50.2 ARTEMISIA VERLOTIORUM 63, S.W. Yorks.: Now extinct at all its known sites.] ©135/50.dra ARTEMISIA DRACUNCULUS __*39, Staffs.: Many plants in pavement, between 43 New Street, Worsley and the Chapel, SO891869, C.B. Westall, 2005. +135/55.4 ANTHEMIS TINCTORIA *94, Banffs.: Three plants growing on a heap of soil on north side of road, Nether Dallachy, NJ362637, I.P. Green, 2005. Several more plants found on 10.9.2005, in same area, some of which were flowering. ©135/56.cor CHRYSANTHEMUM CORONARIUM *71, Man: On a levelled and capped tip at Pooildooie, Ramsey, SC444949, M.M. Davidson & L.S. Garrad, 2000. ©135/61.1 COTULA CORONOPIFOLIA *57, Derbys.: Bed of dried up pond between Ilkeston and West Hallam, SK452415, A. Willmot, 2004. £135/62.1 SENECIO CINERARIA *94, Banffs.: 1 clump in area of waste ground (probably cleared house site), Portsoy, Institute Street, NJ590660, A.G. Amphlett, 2005. 135/62.1x10 SENECIO XALBESCENS (SENECIO CINERARIA X S. JACOBAEA) *71, Man: Ramsey, SC449942, L.S. Garrad, 1997. *H12, Co. Wexford: Three plants amongst rocks on quay, Kilmore Quay, S96-03-, P.R. Green, 2005. 135/62.13x15 SENECIO XBAXTERI (SENECIO SQUALIDUS x S. VULGARIS) *99_ Dunbarton: Waste ground, Craigendoran, NS310816, A. McG. Stirling, 1978. The plant subsequently disappeared but was also found on waste ground, Kilbowie Road, Glasgow (NS4972) by A. McG. Stirling in 1982. ©135/74.1 AMBROSIA ARTEMISIIFOLIA *94, Banffs.: Two plants in flower bed in the garden of 19 Bogmoor Road, Bogmoor. NJ356629, I.P. Green, 2004. *H6, Co. Waterford: Single plant on waste ground, Tramore, $59390219, P.R. Green, 2005, herb. P.R. Green. ©135/79.1 HELIANTHUS ANNUUS *H6, Co. Waterford: Single plant on waste ground, Tramore, $59250235, P.R. Green, 2005. £135/80.1 GALINSOGA PARVIFLORA *77, Lanarks.: By lamp posts and base of wall, Maryhill, Glasgow, NS5668, S.J. Longrigg, 2003, herb. P.M. Reappeared in 2004 and still present 2005. +135/83.1 COSMOS BIPINNATUS *94, Banffs.: One plant in landfill site, Nether Dallachy, NJ365643, I.P. Green, 2005. +135/AGE.hou AGERATUM HOUSTONIANUM *@©63, S.W. Yorks.: Waste ground, Abbeyfield Park, Sheffield, SK3589, Sheffield Wildlife Trust, 2003. +135/CAT.cae CATANANCHE CAERULEA *71, Man: Gravel driveway entrance in Bride Road, Ramsey, SC446954, L.S. Garrad, 2000. ©135/GUL.aby GUIZOTIA ABYSSINICA *85, Fife: Garden, Southerton, Kirkcaldy, NT265911, G. Walker, 2004. Bird seed alien. *94, Banffs.: Several plants in landfill site, Nether Dallachy, NJ365643, I.P. Green, 2004. *135/SIL.per SILPHIUM PERFOLIATUM *71, Man, Bird seed casual in garden, Snugborough, SC355777, M.M. Davidson, 1996. © 137/4.1 ALISMA PLANTAGO-AQUATICA 94, Banffs.: Several plants in pond, Nether Dallachy Landfill Site, NJ364641, I.P. Green, 2005. 1“ record since 1970. *138/4.2 ELODEA NUTTALLI *94, Banffs.: Growing in Pottie Burn just south of track, Bogmoor, NJ354632, I.P. Green, 2004. 151/1.13x14 JUNCUS xSURREJANUS (JUNCUS ARTICULATUS x J. ACUTIFLORUS) 37, Worcs.: Castlemorton Common, SO778398, C.A. Stace & R. Maskew, 2004. Ist record since 1909. £151/1.28 JUNCUS PALLIDUS 37, Worcs.: Lake margin, Lower Moor Gravel Pit, SO0981467, J.J. Day, 1978, det. C.A. Stace. Ist record since 1970. Still present in 1989, A.W.Reid. 151/2.6c LUZULA MULTIFLORA subsp. HIBERNICA *H12, Co. Wexford: Forest ride, Corrageen, S802415, P.R. Green, 2005, det. T.C.G. Rich. 152/13.1 RHYNCHOSPORA ALBA *H6, Co. Waterford: Very plentiful over bog, Bohadoon, S24390265, P.R. Green, 2005, herb. P.R. Green. 152/16.1x15 CAREX XBOENNINGHAUSIANA (CAREX PANICULATA Xx C. REMOTA) *H6, Co. Waterford: Two clumps in damp woodland with both parents, Tournore, X27599587, P.R. Green, 2005. 152/16.44x46 CAREX XFULVA (CAREX HOSTIANA X C. VIRIDULA) *63, S.W. Yorks.: Upland flush, Agden Bog, SK251932, J.G. Hodgson, 1977. PLANT RECORDS 193 152/16.46c CAREX VIRIDULA subsp. VIRIDULA 37, Worcs.: Margin of seasonal pond, Burbury Brickworks, SP100834, I.C. Trueman, 1995, conf. A.C. Jermy. Ist record since 1970. Still present 2004, J.J.Day. +153/10.1 ORYZOPSIS MILIACEA ©63,S.W. Yorks.: Wrose, Shipley, SE157372, B.A. Tregale, 2004. Ist record since 1970. 153/12.4 FESTUCA ALTISSIMA ®96, Easterness: Vegetated bouldery slope above stream, Inverfarigaig, NH537228, G. Jones, 2004. Found in course of SNH survey. 153/13.1x2 LOLIUM XBOUCHEANUM (LOLIUM PERENNE xX L. MULTIFLORUM) *Q4° Banffs.: Growing in field on old airfield, Nether Dallachy, NJ358634, I.P. Green, 2004. £153/17.3 BRIZA MAXIMA *©H6, Co. Waterford: Scattered along roadside, Knockboy, S$63070962, P.R. Green, 2005, herb. P.R. Green. 153/18.9 POA COMPRESSA 94, Banffs.: Growing on the north verge of the road, near the entrance to Nether Dallachy Landfill Site, NJ367640, I.P. Green, 2005. Ist record since 1970. On 23/10/05 found at two locations on disused airfield NJ3663. *H6, Co. Waterford: Top of churchyard wall, Fews, S37420784, P.R. Green, 2005. +153/29.1 GAUDINIA FRAGILIS *34, W. Gloucs.: Semi-improved grassland cut for hay, south of Bishop’s Hill Wood, Wickway, ST733872, J.P. Martin, 2005. *©37, Worcs.: Single large clump in set aside, Lower Howsell, Malvern, SO791488, J.J. Day, 2005, conf. S.J. Leach. +£153/30.1b TRISETUM FLAVESCENS subsp. PURPURASCENS *37, Worcs.: Grass verge, Hawcross, SO761302, K. Barnett, 2000, conf. R. Maskew & W.A. Thompson. £153/38.5 PHALARIS PARADOXA *@©S57, Derbys.: Disturbed ground at entrance to West Park Cemetery, SK487337, R.D. Martin, 2005, det. T.B. Ryves. 153/40.2 CALAMAGROSTIS CANESCENS 13, W. Sussex: Wet woodland, The Moor, Burton Pond, SU974173, N. Sanderson & B. Middleton, 2003, conf. F.L. Abraham. Ist record since 1970. *153/46.2 POLYPOGON VIRIDIS ©63, S.W. Yorks.: Leeds Road, Bradford, SE181350, B.A. Tregale, 2005. Ist record since 1970. 153/47.4 ALOPECURUS AEQUALIS *£94, Banffs.: One plant in flower bed in the garden of 19 Bogmoor Road, Bogmoor, NJ356629, I.P. Green, 2004. Accidental introduction from Christie’s garden centre, Fochabers (v.c.95) where naturalised. +153/47.6 ALOPECURUS MYOSUROIDES *©H6, Co. Waterford: Single plant on road verge next to newly sown lawn, Knockanpower, S214018, P.R. Green, 2005. 7153/50.7 BROMUS SECALINUS _ 85, Fife: Waste ground near a grain store, Kirkcaldy Harbour, NT286921, G.H. Ballantyne, 2005, herb. G.H.B. 1st record since 1919. Imported with grain. 153/51.2 BROMOPSIS BENEKENII *63, S.W. Yorks.: Loversall Delph, SE591004, D. Bailey, 2002. £153/53.1 CERATOCHLOA CARINATA *©63, S.W. Yorks.: Fartown, Huddersfield, SE1418, M.J. Lucas, 2005. *£153/59.3 HORDEUM JUBATUM *94, Banffs.: Growing on the verge of the A96, Newtack, NJ440467, LP. Green, 2004. ©153/60.1 TRITICUM AESTIVUM *94, Banffs.: Several plants in landfill site, Nether Dallachy, NJ367642, I.P. Green, 2005. +153/67.pat SPARTINA PATENS *13, W. Sussex: Single patch c. 10m x 1m at top of saltmarsh, S of West Thorney, SU769017, A. Spiers, 2005, det. E.J. Clement. @ Apparently Ist record for the British Isles. £153/ANE.les ANEMANTHELE LESSONIANA *3, S. Devon: Self-seeding and established, Crediton churchyard, SS8300, L.J. Margetts & L.M. Spalton, 1996, conf. R. Payne & T.B. Ryves. @ £153/FAR.spa FARGESIA SPATHACEA *80, Roxburghs.: Woodland by burn, Harden House, Borthwick Water, NT447149, R.W.M. Corner, 2005, det. C. Stapleton, herb. R.W.M.C. First noted here in 1978. Long established from planting. ©153/PAN.cap PANICUM CAPILLARE *57, Derbys.: Open disturbed ground, Heeley, E of River Sheaf, SK3484, K. Balkow, 2003. Mostly imported soil associated with building new bridge and other developments 158/7.1 COLCHICUM AUTUMNALE *+94, Banffs.: Woodland, Fife Keith Wood, NJ4150, I.P. Green, 2004. 194 PLANT RECORDS *158/10.sax TULIPA SAXATILIS *71, Man: Garden discard, Ballagarey, SC341789, L.S. Garrad, 1998. © £158/12.2 LILIUM PYRENAICUM *H12, Co. Wexford: Two flowering plants on verge, Rathphaudin, $83953544, P.R. Green, 2005. £158/19.1 SCILLA BIFOLIA “*94, Banffs.: Single patch in flower in grassy area between minor road and R. Avon, west side of river, more or less opposite house, NJ154264, A.G. Amphlett, 2004, det. I.P. Green. +158/19.3 SCILLA SIBERICA *94, Banffs.: Three plants, only two flowering, growing on bank on north side of path, just west of bridge, Buckie, NJ419654, I.P. Green, 2004. £158/19.5 SCILLA LILIO-HYACINTHUS *94, Banffs.: Patch at bottom of road bank, north side, Boat o’ Brig, NJ319518, I.P. Green, 2000. +158/21.1 HYACINTHUS ORIENTALIS *94, Banffs.: Growing on heaps of soil by Buckpool tip, NJ409643, I.P. Green, 2005. 158/22.1 CHIONODOXA FORBESII *94, Banffs.: Garden refuse by disused railway line and several small patches on bank below houses, Speyside Way, Aberlour, NJ269432, A.G. Amphlett, 2004. £158/22.luc CHIONODOXA LUCILIAE *94, Banffs.: Growing on two heaps of dumped soil on old airfield, Bogmoor, NJ358631, I.P. Green, 2004. Flowering nicely. First seen 9/4/04. +158/23.lat MUSCARI LATIFOLIUM *£94, Banffs.: A single plant in woodland, by path, at top of flight of steps, close to minor road from village to train station, at junction on SE side of R. Isla, Drummuir, NJ377441, A.G. Amphlett, 2004, det. LP. Green. @ Apparently Ist record for the British Isles. 158/24.6 ALLIUM MOLY *©57, Derbys.: One plant amongst rubbish dumped in old gravel pit in open disturbed ground, Derby, SK362359, A. Mylward, 2004, det. A. Willmot. ©158/24.por ALLIUM PORRUM *H6, Co. Waterford: Clump on road verge, Boheravaghera, X125980, P.R. Green, 2005. £158/24.sat ALLIUM SATIVUM *@©63, S.W. Yorks.: Nailers Rough, Bradford, SE121358, B.A. Tregale, 2005. £158/27.1a AGAPANTHUS PRAECOX subsp. ORIENTALIS *©H6, Co. Waterford: Single plant self-sown in crack in pavement, Dunmore East, S689000, P.R. Green, 2005, herb. P.R. Green. Parent plant in neighbouring garden. [158/31.2 LEUCOJUM VERNUM 37, Worcs.: Error in VCCC; sole record, in Amphlett & Rea 1909, is a transcription error. Delete. ] £158/32.1x3 GALANTHUS NIVALIS x G. PLICATUS *H6, Co. Waterford: Two clumps on road verge, Castlerichard, X00589764, P.R. Green, 2005. 158/35.1 RUSCUS ACULEATUS 94, Banffs.: Large patch in woodland south of cemetery, Bellie Hill, NJ354606, IP. Green, 2005. 1st record since 1970. +158/36.1 ALSTROEMERIA AUREA *94, Banffs.: Two patches, 2.5 x 1.5m and 3 x 2m, in ungrazed area above sea cliff between minor road and coast, west of Whitehills, NJ650654 & NJ649654, A.G. Amphlett, 2005. *158/ASH.lut ASPHODELINE LUTEA *3, S. Devon: Garden escape, Exmouth, SY0080, A. Newton, 1999. @ [£158/ERI.den ERYTHRONIUM DENS-CANIS 85, Fife: Misidentification. Delete VCCC record. | *158/ERI.rev ERYTHRONIUM REVOLUTUM *85, Fife: Now thousands of plants long natur- alised in estate woodland, Wemyss Castle grounds, NT327951, G.H. Ballantyne, 1965. In April 2005 it was learnt (from owner of estate) that the plants were Erythronium revolutum, not E. dens- canis, as previously thought. See BSBI Scottish Newsletter no.28 pages 31 & 32.@ Apparently Ist record for the British Isles. ¢159/2.2 SISYRINCHIUM MONTANUM ~~ *@©57, Derbys.: One clump in disturbed ground on old quarrying/mining site, near Bradwell, SK160801, A. Beck, 2005, det. K. Balkow. +159/2.5 SISYRINCHIUM STRIATUM *@©57, Derbys.: Amongst rubbish dumped in old gravel pit in open disturbed ground, Derby, SK362359, A. Mylward, 2004, det. A. Willmot. +159/5.4 IRIS VERSICOLOR *©63,S.W. Yorks.: Upper Common, Broomhead Moor, Sheffield, SK2296, J. Middleton, 2005. £159/8.1 CROCUS VERNUS *94, Banffs.: Single clump on road verge, Drybridge, NJ4362, A.G. Amphlett, 2004. PLANT RECORDS 195 £159/8.2 CROCUS TOMMASINIANUS *4, N. Devon: Extensively naturalised near a garden, Middleborough, SS6833, N.F. Stewart, 2003. *94, Banffs.: Verge of A97, Aberchirder, NJ627524, A.G. Amphlett, 2004. +159/8.angxfla CROCUS XSTELLARIS (CROCUS ANGUSTIFOLIS x C. FLAVUS) *94, Banffs.: Verge of A97, Aberchirder, NJ627524, A.G. Amphlett, 2004. +159/SCH.coc SCHIZOSTYLIS COCCINEA *94, Banffs.: 2 plants on edge of trees, east side of B9104, Bogmoor, NJ356632, I.P. Green, 2005. +160/3.1 CORDYLINE AUSTRALIS *H12, Co. Wexford: One tree self-sown in pavement at base of wall in Michael Street, New Ross, S71902737, P.R. Green, 2005. 162/8.1 Goodyera repens @®96, Easterness: On slope above Ist bend on path to Falls, Foyers, NH498204, I.P. & P.R. Green, 2002. 162/10.1 HAMMARBYA PALUDOSA ®96, Easterness: 10 plants in area 2m x 3m in hillside flush, S.E. slopes of Meallfuarmhonaidh, NH461220, C. Geddes, 2005. 162/11.1 CORALLORHIZA TRIFIDA ®@85, Kinross: Swampy woodland near edge of loch, Levenmouth Plantation, Loch Leven NNR, NO170005, L. Farrell, 2005. Ist record for hectad since 1920. 162/SER.lin SERAPIAS LINGUA __*3, S. Devon: Single flowering plant in a hay meadow near the coast, Kingsbridge area, SX74, L.M. Spalton, 1998. See BSBI News 79 page 20. © | | ' | | r Bill b f ; Pyne | ih! i a ' Va age ols al i} ill i) 7 a ay uy / i] | f : HTT } il aT ' ean 7 icy Ht | ieee. i} Hill } AN . | if : Wii) JS i} | ee 1 Ni i] | i} i} {i} TA ih Ht is nt Ml | , i | lf i 1) Hi \. i} Hy | itt : j \| \ v i } J i | ay flit l} a { 4 \anhatl il] i} | | | | | : Nie WOH IE | | II / PCM I ul \\ | } a WAHT | Belt | I} bie] fl | WP | 1 Why | } | i} Hh | i} | va va | . J ht | i it = 1 | i } | | | } | | 3 } | hi) i| i | | Hi |} Ht } | ) | ’ | | } | Hel \ HH My {|} Hilt | 1} a , WATMDA INT Wilt Mik | Hitt ait AW yet Wh i} Vy | f an HHT | I} i} | | | | | HUH | i hilt | | Milt} Wid | y) MA oat | I a Mi Wt . i a lal | } | B | Ha Hl if . i % val ane AME rss uh Watsonia 26: 197-211 (2006) 197 Obituaries FRANKLYN HUGH PERRING (1927-2003) Dr Frank Perring, who died of cancer on 11 October 2003, did as much as any other individual to influence the activities of the B.S.B.I. in the second half of the twentieth century. As a young man his partnership with Max Walters ensured the successful completion of the Atlas of the British Flora (1962). This initiated a new era in natural history in the British Isles and much of northern Europe, popularising a method of displaying the results of recording which was to become almost universal at both national and local level. Subsequently he led the Society into a close partnership with the Nature Conservancy and its successor bodies with the recording of rare species for the first British Red Data Book (Perring & Farrell 1977, 1983), before playing a key role in developing the modern Wildlife Trust movement. EARLY YEARS Franklyn was born at Forest Gate, East London, on | August 1927, the son of Frank Arthur (an antiques dealer) and Avelyn Millicent Perring. His early years were spent at Woodford Green at the edge of Epping Forest and at a weekend cottage on the Blackwater estuary. Here, encouraged by his Uncle Stanley, a biology teacher, he became interested in natural history, climbing ancient trees, catching crabs, roach and perch and collecting wild flowers and frogspawn. He received a further stimulus from C. J. Sims, his biology master at Earls Colne Grammar School. After National Service in the army in Northern Ireland, India and Malaya he read Natural Sciences at Queens’ College, Cambridge, obtaining his B.A. degree in 1951 with a 21 in Botany. He went to University thinking that he wanted to be a plant pathologist, but “six weeks in the long vacation in a Pathology Laboratory cured me of that”. In 1949 Frank and his first-year undergraduate friend Richard West decided that they would like to find somewhere in Britain to carry out a botanical survey in the summer vacation where nobody had ever done one. Their enquiries in the Outer Hebrides led to a letter from Peter Morrison of Heisgeir (the Monach Isles) which ended “Come on, boys!” Thus encouraged, they hitch-hiked in a lorry to Glasgow and after two whole days arrived at Paible, from which they reached Heisgeir in a lobster-boat. They used the deserted school- house on Ceann Ear, built in the 1880s, as a base for their studies, returning the following year to complete the work, when they were joined by Michael Locke, a _ like-minded zoology student. Recognising the potential of the building as a base for fieldwork, in 1953 they wrote to Inverness County Council offering to buy it; they received a favourable reply and gave £50 for it, but “got a bit of a shock when £25 was added for legal fees”. In 1966 the Monach Isles were declared a National Nature Reserve under agreements with North Uist Estate, the Commissioners for Northern Lighthouses and the three owners of the schoolhouse; subsequently their European importance has been recognised. Frank and Richard returned to Heisgeir in 1970 with Roland Randall, a visit that led to a Flora of the islands (Perring & Randall 1972). In 1996 Scottish Natural Heritage took a lease on the schoolhouse and in 2004 a rescue plan was launched, leading to several successful work- weeks. When a charitable Trust for Heisgeir was formed in 2004, together with the Friends of the Schoolhouse (Cairdean an Taigh-Sgoile), the owners gifted the schoolhouse to it. Frank stayed in Cambridge to obtain a PhD under the supervision of Dr R. E. Hughes at the Department of Agriculture for his thesis Studies in some chalk grassland plant communities with particular reference to climatic and edaphic gradients (1956a). This led to three papers in the Journal of Ecology (Perring 1958a, 1959, 1960a). When Jon Bennie (2003) repeated Frank’s observations 50 years later he commented that his “detailed, legible notes put my rain-smudged notebooks to shame”. During his undergraduate and postgraduate years Frank’s sense of humour was displayed by the practical jokes he played on the staff, and his editorship of the occasional publication Tea Phytologist. 198 FIGURE 1. Frank Perring at work in the Cambridge office of the B.S.B.I. Maps Scheme. Note the pile of ‘bus tickets’ on the desk in front of him and the row of county floras behind. ATLAS OF THE BRITISH FLORA In 1953 Max Walters asked Frank to become ‘Senior Worker’ for the B.S.B.I. Distribution Maps Scheme’, a post he accepted and took up in October 1954. It appears from Frank’s account (Perring 1992) that Max “popped the question” on a botanical excursion to Odsey; there is no suggestion that the job was advertised or that other applications were considered. However unorthodox these recruit- ment procedures may appear to a _ later generation, this was a job ideally suited to Frank’s talents and the one in which he was to make his name. With boundless energy, much charm and a character which was completely impervious to discouragement, Frank set about mobilising the membership of the B.S.B.I., and the wider public, to record the plants of some 3,500 10-km grid squares in Britain and Ireland. The Society, described by Max Walters (1954) at the start of the scheme as “flourishing and expanding”, was in a good position to take on the project, but it involved a considerable departure from its traditional practices. Senior figures, long set in their ways and with little experience of co-operative research, had to be persuaded to adopt the new taxonomy of ‘CTW’ (first published in 1952), trained to read Ordnance Survey grid references and encouraged to use the unfamiliar field cards (hitherto used in only a few county flora schemes). Frank’s first lecture on the scheme, he once related, was in North-East England, to an audience which included the formidable | OBITUARIES Professor J. W. Heslop Harrison. Frank succeeded in recruiting even him to the cause, and Heslop Harrison was to contribute master cards to the scheme for the Hebridean islands, restricting his objections to occasional taunts in the pages of the Vasculum. B.S.B.I. field meetings were arranged to visit areas where resident botanists were absent, and a generation familiar with National Service soon came to refer to the new recording methods as ‘square- bashing’. New recruits were sought from the wider public: a single article in The Observer by John Gilmour brought over 800 replies. In the Republic of Ireland, where manpower was particularly short, David Webb (sometimes accompanied by Frank) played a crucial role in field recording and data extraction. In addition to field work, the other aspect of data-gathering was the collection of records of rarer species from herbarium specimens and _ literature ~ sources. In this too Frank played a major role, extracting many thousands of records onto 40- column ‘individual record cards’, colloquially known as ‘bus tickets’ (Fig. 1). The extent to which the scheme drew on all available recorders is illustrated by the concessions made to less expert recorders. A special card was prepared for those unwilling to relinquish the taxonomy of Bentham & Hooker: those species which were too broadly defined to be meaningful in the 1950s were listed but lacked code numbers by the name, so that they would not be entered onto punched cards. Another card, specially designed for use by school children, listed the scientific and English names of 113 common and supposedly unmistakable species. A key part of the project was the use of mechanised data-handling methods. Field card records were transferred to “master cards’ and then each one punched individually onto the 40-column cards, with holes denoting species code number, vice-county, grid reference and date. The cards (together with the individual record cards) could then be sorted into species and grid square order. They were then fed through a tabulator for mechanical mapping by a specially modified (and, at the time, unique) machine (Fig. 2). As so often happens, the time taken to process the records was _ under- estimated, not least because the success of the scheme had brought in more data than expected. The initial plan to complete data collection in 1959 and publish in 1960 proved impossible and, “to the consternation of the Nature Conservancy”, who, with the Nuffield ' Walters (1954), Walters & Perring (1956), Perring & Walters (1962), Perring & Sell (1968), Allen (1986), Harding & Sheail (1992) and Perring (1992) describe the scheme from various perspectives. OBITUARIES 1199 FIGURE 2. Frank Perring demonstrating the tabulator modified to print the maps for the Atlas of the British Flora. Foundation, had financed the scheme, two extensions of the original deadline were required (Harding & Sheail 1992). The project was successfully brought to completion with the publication of the Atlas of the British Flora in 1962. Publication was celebrated at a special dinner in Chelsea on 28 April. It was on this occasion that Frank produced what was perhaps his most celebrated pun when, in paying tribute to Max Walters, he referred to “the most important factor in the Scheme — the ‘Max’ Factor’. In his speech Lord Hurcombe said that “although the Nature Conservancy had spent £25,000 financing the work they felt they now had good value for money’. He drew attention to the potential use of the Atlas in preventing the loss of treasured national resources, an early appreciation of its potential use in conservation work. Max Nicholson commented that “ornithologists ... had been put to shame by the botanists, and other Societies would soon have to follow the lead given by the B.S.B.I.” (Bowen 1962). These were perceptive words, and the Aflas proved to be one of the most influential natural history books published in the 20th century. OTHER WORK AT CAMBRIDGE 1954-1964 While taking part in the more general work of the Botany Department, Frank became in- terested in Arctium. With Peter Sell, he grew a large range of Arctium plants in an experiment which covered half an acre of experimental plots in the University Botanic Garden (Sell 2004). This led to an early piece of ‘network research’ in which B.S.B.I. members were invited to measure plants in the field (Perring 1960b), to Frank’s account of Arctium in Flora Europaea (1976a), his one formal contribution to that major collaborative project, and eventually to the taxonomic conclusions summarised in the recent volume of Sell & Murrell’s (2006) Flora. In tandem with work for the Atlas, during the 1950s Frank gave special attention to botanical records for Cambridgeshire (v.c. 29). Max Walters was B.S.B.I. vice-county recorder from 1949 to 1961, then handed over to Frank, who served until 1971. They worked on the vice- county records with Peter Sell, publishing lists in the B.S.B.I.’s Proceedings and in Nature in 200 Cambridgeshire (e.g. Perring, Sell & Walters 1955, Perring 1956b, 1958b, Perring & Sell 1959) and planning for a new county flora in 1960 to commemorate the tercentenary of John Ray’s Catalogus (the very first County Flora) and the centenary of C. C. Babington’s Flora of Cambridgeshire. This was to be based on the Cambridge Natural History Society’s card index and site lists, from which and from records submitted for the Atlas Frank compiled a “master “hist In! the event Ay Hlora of Cambridgeshire appeared in 1964, with a section on Bryophyta by Harold Whitehouse, and was the first County Flora for vascular plants to use O.S. 10-km grid squares (though not as yet dot maps) to indicate plant distributions. CHARACTER AND INTERESTS Frank was irrepressibly exuberant, with an infectious enthusiasm and a_ seemingly inexhaustible fund of ideas (as well as puns, some brilliant, some execrable). He made no real distinction between botany as his pro- fession and as a spare-time passion, whether in fieldwork or when lecturing, teaching and writing. “One of Frank’s great assets was his combination of a businesslike and at the same time cheerful manner. Bustle, in the best possible sense of the use of the word, I would always associate with him, and he never allowed rules and regulations to get in his way (Sell 2004). He would indeed head straight towards his goal, oblivious of the fact that he was cutting across official pathways. When faced with an opposing argument he was never at a loss for a highly plausible response. He was not so much intolerant of opposition as incredulous that it could exist; criticism rolled off him, and he never bore a grudge. Nevertheless, those with more conventional minds could sometimes find him = an exasperating colleague, and not all of them shared his forgiving nature. He was at his best with volunteers, whereas his methods could put considerable strain on those who worked under him. Few failed as spectacularly to cope with him as the Monks Wood secretary who, on her first day at work, left in tears at lunchtime, never to be seen again. In an unpublished ‘Biographical Sketch’ written in 1965, Frank summed up his non- botanical interests: “Rarely if ever relax — read perhaps two novels a year — at Christmas and during’ the summer holiday -— nearly always spent abroad ... Favourite reading Jane Austen and OBITUARIES Dickens: take one of the latter on each long journey: I remember reading Pickwick in the Pickwick Hotel, San Francisco. Very fond of music, especially Mozart and Britten. Britten’s Spring Symphony and Elgar’s Introduction and Allegro and Violin Concerto spell out for me the Englishness of England which I want to see preserved.” To these could be added some equally English but less highbrow interests, including West Ham football club, rugby, horseracing (but not betting) and bell-ringing. And England was not his only love: he said earlier in the same Sketch that his travelling on field meetings had made him “a strong ‘Britainophil’, particularly fond of Ireland which I try to visit at least once a year — just to enjoy the slower pace and the brighter talk”’. HEAD OF THE BIOLOGICAL RECORDS CENTRE 1964-1978 Originally, it was planned that, at the completion of the Atlas project, the Nature Conservancy would take over the machinery and record cards “as the basis of a permanent recording system” (Walters 1954). In 1963, the Nature Conservancy agreed to transfer Frank and his staff to the recently opened Monks Wood Experimental Station near Huntingdon. (Frank had become Director of the Distribution Maps Scheme in 1959 when Max Walters resigned after the first five years of the project.) Throughout his period at Monks Wood Frank served on the. B.S.B.I. Maps Committee, which was renamed Records Committee in 1966, and he was Secretary of the Records Committee from 1967 to 1981. These twin positions gave him a dominating influence on the recording activities of the Society. As Head of B.R.C., Frank’s job was to continue the work on vascular plants initiated by the Atlas project and to set up similar schemes for other groups of organisms. The Critical Supplement to the Atlas of the British Flora (Perring & Sell 1968) dealt with several major critical genera, and included complete treatments of Alchemilla, Euphrasia, Hieracium and Sorbus and numerous other critical species, hybrids and infraspecific taxa. In the Critical Supplement the maps are accompanied by explanatory text, and the volume remains a valuable work of reference. It was another product of Frank’s productive partnership with Peter Sell, and the accounts of Hieracium and Pilosella, contributed by Sell and West, occupied over a third of the book. OBITUARIES Frank had intended to round off the Atlas work with a book on the phytogeography of the British flora, and (with Max Walters) he had actually signed a contract to write one. However, even before the publication of the Critical Supplement his primary interests had switched to plant conservation, which was henceforth to be his major botanical pre- occupation. The maps in the Atlas had provided a very dramatic visual representation of the rarity of some species and the decline of others. Frank decided to focus B.R.C.’s botanical work on gathering more precise data on _ the distribution of the rarest species, and Miss M. N. Hamilton was recruited as a new member of staff to carry out this work. In 1968-69 lists of the known localities of species believed to be confined to 1-8 and 9-15 10-km squares were circulated to B.S.B.I. vice-county recorders, Nature Conservancy staff and others, with a request for updated information on each population. Additional impetus was given to the work by the Wild Plants Protection Bill, a Private Member’s Bill introduced in 1968 by the Conservative MP Peter Mills which failed to pass into law but nevertheless highlighted the need for information on the rarest species. The work was widely publicised (e.g. Perring 1971a, Perring & Walters 1971) and eventually led to the pioneer British Red Data Books: 1 Vascular Plants (1977), compiled by Frank and Lynne Farrell, Noelle Hamilton’s successor at B.R.C. Frank’s other initiatives at B.R.C. included ‘network research’ projects. The most successful of these was the Winter Mistletoe Count, a survey conducted in the winters of 1969/70 to 1972/73 to map the distribution of Viscum album at tetrad scale (Perring 1973). By 1975 four Network projects were underway, recording arable weeds, black poplars (Populus nigra), Symphytum taxa and holly (lex aquifolium) (see Watsonia 10: 294-299, 1975). The investigations of the taxonomy and distribution of Symphytum arose from the confusion of S. officinale and S$. xuplandicum in the Atlas survey (Perring 1969a, 1970a, 1975). Frank took a particular interest in the genus, eventually writing, jointly with Professor Theo Gadella, the account in The European Garden Flora (Gadella & Perring 2000). The early 1970s were a productive period in Frank’s career. He edited The Flora of a changing Britain (Perring 1970c) and The British Oak (Morris & Perring 1974) with an entomological colleague at Monks Wood. This book was based on papers given at a highly 201 successful B.S.B.I. conference held at the University of Sussex in 1973 and achieved a wide circulation. It was often cited as a recom- mended text for University courses, not only in Britain but even as far afield as Iceland (where there are no oaks). His contribution (1974) to The changing flora and fauna of Britain Symposium made effective use of distribution maps to illustrate “changes influenced by man” and “changes not influenced by man”, as did several other papers in the same volume. Also published in 1974 was English names of wild flowers (Dony, Perring & Rob 1974) which provided standard English names by using hyphenation to generate a series of English ‘genera’ and ‘species’. A standard list was required as ‘CTW’ did not provide English names for all species, but the subject proved to be remarkably emotive and the treatment developed for this slim volume continues to excite controversy. Frank played a key part in launching Atlas Florae Europaeae. In this project the taxonomic work of Flora Europaea (1964— 1980), edited by a team of British botanists including Max Walters, was united to the methods developed by Frank and Max for the Atlas, with the additional inspiration of Hultén’s earlier Atlas 6ver Vdxternas utbredning i Norden (Perring 1967a). As the first volume of Atlas Florae Europaeae stated, “a map presented by Dr. Perring at the Tenth International Botanical Congress in Edinburgh, 1964, which showed the 50-km_ square distribution of Silene acaulis in Europe, can be regarded as the starting-point of the present work” (Jalas & Suominen 1972). Frank acted as secretary to the nascent committee, visiting a number of European countries in 1965 and producing a further nine maps at Monks Wood before responsibility for the Secretariat was handed over to Helsinki in December 1965. During his period at B.R.C. he was the British regional collaborator on the Committee for the Mapping of the Flora of Europe, and he remained an active Advisor to the Committee until his death. In addition to the formal recording initiatives he launched at Monks Wood, Frank gave generously of his time to inexperienced botanists, both young and old. The revival of the county flora tradition in the 1960s owed much not only to the recording methods pioneered by the Atlas team but also to the personal encouragement he offered to prospective authors. During 1961 and 1962 he was Secretary of a B.S.B.I. ad hoc committee set up, after a conference on Local Floras, to 202 draw up guidance for Flora writers; the results, including a list of recent and forthcoming Floras, were published in Proceedings in 1964 and revised in 1971 (Perring 1964, 1971b). In the preface to his Flora of Flintshire, Goronwy Wynne (1993) has recounted what happened when he visited Monks Wood to ask for Frank’s advice on what he should do as vice- county recorder. “He suggested in no uncertain terms that it was high time that Flintshire had a county Flora of its own, and that I, as Recorder, should take it on. I left that meeting with a mixture of euphoria and sheer terror ... .””- Frank also ran many field courses, both for the Field Studies Council (especially at Preston Montford) and the Shropshire Adult College at Attingham Park in Shropshire, one of the counties he ‘adopted’. Four of the latter courses, held between 1969 and 1972, led to the production of a duplicated Flora of Attingham in 1975. (Sarah Whild and Alex Lockton have recently published a new Flora based on a comparison of the current flora with that documented by Frank’s teams.) Ten years later Ecological Flora of the Shropshire Region was published (Sinker et al. 1985). Frank was author of only one chapter, on biogeographical elements, but his influence in the preparation and recording for the project was much greater than this might suggest. As Head of B.R.C., Frank worked tirelessly to promote biological recording on numerous journeys abroad and in international and national journals (e.g. Perring 1967b, 1969b, 1970b, 1971c, 1972, 1976b, c, Soper & Perring 1967, Heath & Perring 1975) as well as in many local publications. He presided over the computerisation of the database in 1970-71, when record cards were transported to London in batches of 300,000 to be transferred to magnetic tape. Thereafter he promoted the role of the Centre as a computer database and proved adept at publicising its activities. On one occasion in 1972, after an appeal for recorders on the television programme Tomorrow’s World, B.R.C. was submerged under an avalanche of mail which it took his staff several weeks to process. The role of B.R.C. in zoological recording expanded after 1967, when John Heath was appointed to the staff. Frank was in the forefront of the campaign to establish Local Records Centres (Perring 1976d), although for many years their development was hampered by uncertain funding. Although Frank’s years at B.R.C. were marked by many _ successes, he was OBITUARIES increasingly hampered by his inability to work with his managers. B.R.C. was run as a self- contained unit which was not well integrated with other sections at Monks Wood. Even in the Nature Conservancy days B.R.C.’s frog emblem (it would now be described as a logo) had come to symbolise Frank’s independence of view, and at one meeting Martin Holdgate, then a Deputy Director, commented that, as far as he was concerned, “the frog could jump into the lake”. Matters became worse after 1973 when the Nature Conservancy was split to form two organisations, the Nature Conservancy Council and the Institute of Terrestrial Ecology. Frank lobbied as hard as he could to get B.R.C. placed in N.C.C., even inspiring the Earl of Cranbrook to enter a plea for this option in a House of Lords debate (12 April 1973). However, there were influential figures in N.C.C. who were determined not to have him in their organisation, and B.R.C. remained at Monks Wood under the new I.T.E. ‘The split’ traumatised Monks Wood, and Frank, in particular, was unable to get on with the rather rigid I.T.E. management. One occasion when the Director, John Jeffers, feigned sleep while Frank addressed a party of visiting dignitaries encapsulated the situation and became in- corporated into Monks Wood’s oral history. It is also true that Frank was a much better tactician than he was _ strategist. Current initiatives brought out the best in him, but the long-term planning needed to run a database was often neglected in favour of the short-term fix. In his later years B.R.C.’s technological capacity actually decreased, as the Atlas methods became obsolete but I.T.E. failed to invest in adequate replacements. By the late 1970s, for example, mechanical mapping was no longer possible and maps were hand-plotted by members of B.R.C. staff. To some extent the true situation was concealed from the outside world by Frank’s presentational skills, although his tendency to over-commit B.R.C. was a further cause of problems, especially as he could not rely on management support in meeting his promises. Frank was never interested in organisational niceties and made no distinction between the activities he undertook for I.T.E. and those which he undertook for B.S.B.I. LT.E. management increasingly tended to regard him as a B.S.B.L. employee whom they had the misfortune to pay. When he left B.R.C. at the end of 1978 it needed all Dick David’s negotiating skills as B.S.B.I. President to ensure that a replacement botanist was appointed to the staff. OBITUARIES BOTANICAL SECRETARY OF THE LINNEAN SOCIETY 1973-1978 Frank was an active Fellow of the Linnean Society of London for 39 years. As Botanical Secretary from 1973 to 1978, he helped to organise a number of successful conferences at Burlington House, including one on bracken, the proceedings of which he edited with Brian Gardiner, the Zoological Secretary, for publication in 1976 (Perring & Gardiner 1976). Professor Irene Manton was then President of the Linnean Society — its first woman President and a forceful character — and she and Frank made a good team. In January 1978 the Society celebrated the bicentenary of Linnaeus’ death and Frank also participated in its visit to Sweden at the end of June that year. In April 2002 he organised a _ well-attended joint Linnean Society/B.S.B.I. meeting entitled Where do we find and train the next generation of field botanists? WORKING AS A VOLUNTEER WITH THE NATURALISTS’ TRUSTS Frank’s involvement with the (then) Naturalists’ Trusts began in 1956, when the Cambridgeshire and Isle of Ely Naturalists’ Trust (CAMBIENT) was set up. He served on its Council continuously until a revision of the constitution brought about his retirement after a quarter of a century in 1982. He was Secretary of the Technical Committee from 1957 till the end of 1967, a post which gave him useful knowledge of the operation of nature conser- vation legislation (particularly the S.S.S.L system) and experience in assessing the value and management requirements of specific sites. He became Secretary of a reformed Field Committee in 1964, a post he held till 1968, when he continued as a committee member. Initially this committee co-ordinated the efforts of 18 voluntary ‘regional officers’ and 25 ‘watchdogs’ appointed to look out for dev- elopments likely to harm sites, and later it took on responsibility also for work parties and field excursions and began the formation of local groups; it thus combined interests which were dear to Frank throughout his life — the harnessing of volunteer labour (for both conservation and recording), and public educ- ation. In his 1965 ‘Sketch’ he said, “I spend most of my time in the winter in practical conservation — leading parties bush clearing, or tree felling in the areas owned or administered by the Trust.” From 1968 till 1970 he was Chairman of the Executive Committee. 203 Frank moved to Oundle in 1971 and the focus of his activities moved away from Cambridgeshire. He was already interested in the botany of Northamptonshire, having produced a) paper for ‘the B:S.Bih‘s Proceedings in 1955 on the plants of the county mentioned in John Clare’s poetry and prose works (Perring 1955). By 1976 he was active in fundraising for the Northamptonshire Trust, organising Medieval Fairs at Southwick in 1977 and 1979 and at Ashton in 1981, to help purchase reserves such as Short Wood, Glapthorn Cow Pastures and Titchmarsh Heronry. He served on the Trust’s Council and Executive Committee and as its Chairman from 1985 to 1987. He was joint editor of The Nature of Northamptonshire (Colston & Perring 1989). In 1987 Frank formed the Peterborough Wildlife Group, becoming its first Chairman. The Northamptonshire Wildlife Trust celebrated its 30th birthday in 1993; in the same year it, the Cambridgeshire Trust, the Beds & Hunts Trust and the P.W.G. united as a single Wildlife Trust and Frank joined the Editorial Board of its new publication, Wildlife Action. GENERAL SECRETARY OF THE ROYAL SOCIETY FOR NATURE CONSERVATION 1979-1987 In 1979, at the age of 51, Frank left Monks Wood to become General Secretary of the Society for the Promotion of Nature Conservation, the umbrella body for 41 autonomous Naturalists’ and Nature Conservation Trusts, which changed its name to Royal Society for Nature Conservation in 1981. Believing passionately in the importance of the Trusts and their work at the local level, he also recognised that the cause of nature conservation had to be promoted more widely and that the Trusts would benefit from more co-ordination and more financial resources. In 1981, against all odds, Frank was able to launch a new national magazine, Natural World, available to all Trust members. He campaigned to standardise the names of the Trusts, suggesting that “Naturalists’ Trust” conjured up a ‘fuddy-duddy’ picture in many people’s minds and even (rather implausibly) that there was a danger of confusion with naturists; he argued forcibly that one name — “Wildlife Trust” — would present the Trusts as a united movement. There was considerable Opposition at the time, and not all the Naturalists’ Trusts had changed their names by 204 OBITUARIES the time Frank retired, but in the end even the first-established Naturalists’ Trust acceded and changed its name to “Norfolk Wildlife Trust” and R.S.N.C. itself has adopted the title “Royal Society of Wildlife Trusts”. In October 1985 the Society launched its British Wildlife Appeal under the patronage of the Prince of Wales and the chairmanship of Sir David Attenborough, raising £15 million for the Trusts by the end of 1990. That the concept of a national appeal was accepted by the Trusts as the way forward reflects considerable credit on Frank’s powers of argument and persuasion and it undoubtedly enhanced their standing. Frank enthusiastically embraced the burgeoning Urban Wildlife Group movement, successfully bringing it into the R.S.N.C. ‘family’ and seeing it as an opportunity to engage with a new and often younger audience. During his eight and a half years at R.S.N.C. the number of constituent Trusts rose to 48, Watch Club (created in 1975 for young people) grew rapidly, some 500 nature reserves were established (to reach a total of 1,680), over 30 Urban Wildlife Groups were founded, and the Trusts’ membership rose from 129,000 to 180,000. As in L.T.E., Frank’s time at R.S.N.C. was not all plain sailing: he got on well with David Attenborough (Fig. 3), but his relations with Walter Lane, R.S.N.C.’s Executive Chairman, were stormy. Nevertheless, his energy, entre- preneurial flair and cheerful optimism usually triumphed and were widely appreciated by his colleagues. He was appointed O.B.E. in 1988 for services to nature conservation and claimed, characteristically, that it must stand for “OBsessive Enthusiasm”. AN ACTIVE RETIREMENT 1987-2003 Frank retired from R.S.N.C. on 31 July 1987 but continued to work tirelessly. An early project was to write, with Max Walters, The Macmillan Field Guide to British Wildflowers (1989), a guide illustrated by Andrew Gagg’s photographs. Frank was primarily responsible for the “apt, pithy and sometimes amusing descriptions [which] are, in fact, the greatest merit of the book” (Oswald 1990), e.g. the “rugby-ball-shaped pods” of scurvygrasses and common whitlow-grass and the “wide-awake looking” flowers of common dog-violet. A less well-known publication was the text he wrote FIGURE 3. David Frank Perring with Sir Attenborough at the President’s Reception, Royal Society for Nature Conservation, November 1993. Photograph: R.S.N.C. (anonymously) for a wild flower poster distri- buted in Woman magazine (14 May 1988); the same issue features “the hottest film heart-throb - in Hollywood”. In 1988 Frank set up, with Anne Cryer, and chaired Wildlife Travel, a company offering expert-led overseas holidays in search of flowers and other wildlife and donating its profits to the Wildlife Trusts (more than £100,000 over 17 years). Frank himself led or co-led some 50 of the trips, first to the Mediterranean but later to more distant places including Australia, Nepal, St Petersburg and the Arctic, China and South Africa. Many of the company’s clientele became ‘regulars’ and the annual get-togethers were gatherings of friends rather than customers. Frank was elected President of the B.S.B.I. at the 1993 A.G.M. at Lincoln, giving his Presidential Address, ‘Druce in Northampton- shire’, the following year (Perring 1995). He was elected an Honorary Member in 1997. The rather belated Presidency perhaps reflected the reluctance of the Society to give such an unpredictable person a position of at least nominal authority, and the Honorary Member- ship was certainly long overdue. But as far as one could tell Frank took no offence and continued his cheerful service to the Society in several capacities. He remained a familiar figure at the Society’s main meetings, and in particular at the Society’s Annual Exhibition Meeting (which he attended for 49 successive years) and the Scottish Exhibition Meeting. When the Society put the sale of its public- ations in the hands of F. & M. Perring as ‘BSBI Books’, most members assumed that Frank was in partnership with his wife Margaret; in fact ‘F.” was Margaret’s father-in- OBITUARIES law Frank, and she fiercely defended her independence in this venture, even after the death of Frank senior in 1982, saying that it was one thing which she was not going to allow her husband to take over! However, he was usually to be seen helping on the stall at these B.S.B.I. meetings. From the time of his PhD study Frank had a special interest in the flora of France. In retirement he took a lead in the B.S.B.I.’s relations with French and other European botanists, taking part in several joint events, culminating in the symposium Botanical Links in the Atlantic Arc held in Camborne, Cornwall, in May 2003; this was principally concerned with studies of plant species and species-assemblages occurring on or close to the Atlantic-facing coasts of Europe and was attended by over a hundred delegates from Ireland, Scotland, Wales, England, France and Sweden. The resulting book (Leach et al. 2006) is dedicated to Frank’s memory; though in failing health, Frank enjoyed the occasion, and he would have enjoyed also the pun in the preface (if indeed he did not originate it): “For a large number of species the Atlantic Arc is clearly an important refuge — not just an ‘arc’, but also an Atlantic Ark.” Over his last ten years Frank led a B.S.B.I. initiative, in collaboration with the Field Studies Council and the University of Birmingham, to help people get to grips with species identi- fication and biological recording. He set up a working group within the B.S.B.I. (a charac- teristic move, allowing him to bypass the Society's committee structure!) to consider training and education and brought together an impressive array of botanical educators from around the country. Frank’s vision was to launch a suite of training courses, backed up by academic accreditation, to provide a rigorous but enjoyable introduction to botanical fieldwork within the context of biological recording. The B.S.B.I. identified a special need among those working in nature conservation, so the main university certificate has been particularly designed for professionals in Wildlife Trusts, consultancies and Government Agencies and those seeking such employment, though keen amateur naturalists are not excluded. Many hundreds of students have now passed through the programme and it is recognised by em- ployers as a course which will give practical and reliable identification skills. Frank believed passionately in ‘catching them young’ (see Perring 1969c) and, with 205 handsome financial donations from the late Allan Hamerschlag, with whom he and Margaret became friends on a Wildlife Travel holiday in Australia, he worked with the F.S.C. to organise “Spotlight on Plants’ courses for sixth-formers at Field Centres. He also collaborated with John Hewitson of Oundle School and ‘Science and Plants for Schools’ and others to produce A Key for Identifying British Trees and Shrubs on the internet (www- saps.plantsci.cam.ac.uk/trees/). This friendship produced spectacular results for the B.S.B.I. after Frank’s death, as in 2005 Allan Hamerschlag left the Society its largest ever legacy. At Frank’s funeral, on 24 October 2003, a very large congregation bade him farewell in St Rumbold’s Church, Stoke Doyle, North- amptonshire, in a way he would have fully appreciated. Frank had been Secretary to the Stoke Doyle Parochial Church Council, and he was instrumental in changing the management of the churchyard so that it became a flowery meadow. At his remarkable memorial service Max Walters spoke of their long friendship: the text of his address is published in BSBI News 95: 66-67 (2004). FRANK’S CONTRIBUTION TO BRITISH AND IRISH BOTANY We hope that this obituary gives some idea of the range of Frank’s activities, though we have certainly not covered all those that are known to us, and of course there must be others of which we are unaware. Although the Atlas of the British Flora was planned before Frank joined the team, his creativity is apparent from his involvement in the British Red Data Books: 1 Vascular Plants and in Atlas Florae Europaeae, and in his role in reforming the Wildlife Trusts. He was sufficiently unorthodox to come up with new ideas and press for _ their implementation, while being just sufficiently orthodox to be able to work with established organisations. A constant theme running through his career was his desire to communicate his love of botany and belief in the necessity of plant conservation outward to new audiences. When harnessed, his energies could produce remarkable results, as the Maps Scheme showed. Subsequently no organisation perhaps succeeded in getting the best out of him, and certainly the tensions within the organisations in which he worked were more apparent. Nevertheless, his achievements were 206 OBITUARIES formidable. With the exception of the nine ACKNOWLEDGMENTS years at R.S.N.C., the B.S.B.I. was central to his activities from the time he was asked to join Me ate ee to ees ae Carmen the ‘Maps Scheme’ in 1953 until his death fifty ana at ae Se S Bs aa an years later. Frank can justly be regarded as one ahha Se a bee ae hae: ee of the B.S.B.I.’s pivotal figures in the post-war ES ane a DENN J enn is period, one of those who did most to change it Goumeny, amino Oa Beis Wine as from the Society left by G. C. Druce when he Botanical Secretary of the Linnean Society is died in 1932 to that which it had become at the ‘Ket! tom! obituaries’ by Johny Eanes end of the millennium. The Times and The Linnean. REFERENCES ALLEN, D. E. (1986). The botanists. St Paul’s Bibliographies, Winchester. BENNIE, J. J. (2003). The ecological effects of slope and aspect in chalk grassland. PhD thesis, University of Durham. BOWEN, H. J. M. (1962). Dinner to celebrate the publication of the “Atlas of the British Flora’. Proceedings of the Botanical Society of the British Isles 4: 490. COLSTON, A. & PERRING, F. [H.] (1989). The Nature of Northamptonshire. Barracuda Books, Buckingham. Dony, J. G., PERRING, F. [H.] & Ros, C. M. (1974). English names of wild flowers. Butterworths, London. GADELLA, T. W. J. & PERRING, F. H. (2000). Symphytum Linnaeus, in CULLEN, J. et al., eds The European Garden Flora Volume VI Dicotyledons (Part IV), pp. 138-141. Cambridge University Press, Cambridge. HARDING, P. T. & SHEAIL, J. (1992). The Biological Records Centre: a pioneer in data gathering and retrieval, in HARDING, P. T., ed. Biological recording of changes in British wildlife, pp. 5-19. 1.T.E. symposium no. 26. HMSO, London. HEATH, J. & PERRING, F. [H.] (1975). Biological recording in Europe. Endeavour 34: 103-108. JALAS, J. & SUOMINEN, J. (1972). Atlas Florae Europaeae 1 Pteridophyta (Psilotaceae to Azollaceae). Committee for Mapping the Flora of Europe and Societas Biologica Fennica Vanamo, Helsinki. LEACH, S. J., PAGE, C. N., PEYTOUREAU, Y. & SANFORD, M. N., eds (2006). Botanical links in the Atlantic Arc. B.S.B.I. Conference Report no. 24. Botanical Society of the British Isles. Morris, M. G. & PERRING, F. H., eds (1974). The British Oak: its history and natural history. B.S.B.1. Conference Report no. 14. E. W. Classey, Faringdon. OSWALD, P. [H.] (1990). Review: The Macmillan Field Guide to British Wildflowers. Nature in Cambridgeshire No. 32: 73-76. PERRING, F. H. (1955). John Clare and Northamptonshire plant records. Proceedings of the Botanical Society of the British Isles 1: 482-489. PERRING, F. H. (1956a). Studies in some chalk grassland plant communities with particular reference to climatic and edaphic gradients. PhD thesis, University of Cambridge. PERRING, F. H. (1956b). An ms. in Wisbech museum. Proceedings of the Botanical Society of the British Isles 2: 133: PERRING, F. [H.] (1958a). A theoretical approach to a study of chalk grassland. Journal of Ecology 46: 665— 679. PERRING, F. H. (1958b). Additions to the vascular flora of Cambridgeshire. Nature in Cambridgeshire No. 1: 27-29. PERRING, F. [H.] (1959). Topographical gradients of chalk grassland. Journal of Ecology 47: 447-481. PERRING, F. [H.] (1960a). Climatic gradients of chalk grassland. Journal of Ecology 48: 415-442. PERRING, F. [H.] (1960b). Report on the survey of Arctium L. agg. in Britain, 1959. Proceedings of the Botanical Society of the British Isles 4: 33-37. PERRING, F. [H.], comp. (1964). A guide for local Flora writers. Proceedings of the Botanical Society of the British Isles 5: 283-302. PERRING, F. H. (1967a). Mapping the flora of Europe. Proceedings of the Botanical Society of the British Isles 6: 354-357. PERRING, F. H. (1967b). The processing of biological data. Institute of Biology Journal 14: 17-19. PERRING, F. [H.] (1969a). Symphytum survey. Proceedings of the Botanical Society of the British Isles 7: 553— 556. PERRING, F. [H.] (1969b). Mapping the flora and fauna of the British Isles. Country-side Spring 1969: 100- 104. PERRING, F. [H.] (1969c). The demand for and supply of field biologists. Journal of Biological Education 3: 123-129. PERRING, F. H. (1970a). The B.S.B.I. Symphytum survey. Watsonia 8: 91. PERRING, F. [H.] (1970b). The British biological recording network, in CUTBILL, J. L., ed. Data Processing in Biology and Geology, pp. 115-121. Systematics Association Special Volume no. 3. Academic Press, London. OBITUARIES 207 PERRING, F. H., ed. (1970c) The Flora of a changing Britain. B.S.B.I1. Conference Report 11. E. W. Classey, Hampton. PERRING, F. H. (1971a). Rare plant recording and conservation in Great Britain. Boissiera 19: 73-79. PERRING, F. H. (1971b). The vice-county guide. Watsonia 8: 435-447. PERRING, F. H. (1971c). The Biological Records Centre — a data centre. Biological Journal of the Linnean Society 3: 237-243. PERRING, F. [H.] (1972). The great wildlife plot. Birds 4: 72-76. PERRING, F. [H.] (1973). Mistletoe, in GREEN, P.S., ed. Plants wild and cultivated, pp. 139-145. E. W. Classey, Hampton. PERRING, F. H. (1974). Changes in our native vascular plant flora, in HAWKSWORTH, D. L., ed. The changing flora and fauna of Britain, pp. 7-25. Academic Press, London. PERRING, F. H. (1975). Symphytum L., in STACE, C. A., ed. Hybridization and the Flora of the British Isles, pp. 353-355. Academic Press, London. PERRING, F. H. (1976a). Arctium L., in TUTIN, T. G. et al., eds Flora Europaea Volume 4 Plantaginaceae to Compositae (and Rubiaceae), p. 215. Cambridge University Press, Cambridge. PERRING, F. H. (1976b). The future and electronic data processing, in SIMMONS, J. B., BEYER, R. I. BRANDHAM, P. E., LUCAS, G. LL. & PERRY, V. T. H., eds Conservation of threatened plants, pp. 113- 117. Plenum Press, New York and London. PERRING, F. [H.] (1976c). Records for leisure and profit. New Scientist 23/30 December 1976: 725-727. PERRING, F. H. (1976d). The value of museums as records centres, in BOYLAN, P. J., ed. Museums Association Annual Conference Bristol 1976, pp. 19-21. Museums Association, London. PERRING, F. H. (1992). BBSI distribution maps scheme — the first 40 years, in HARDING, P. T., ed. Biological recording of changes in British wildlife, pp. 1-4. 1.T.E. symposium no. 26. HMSO, London. PERRING, F. [H.] (1995). Presidential Address, 1994. Druce in Northamptonshire. Watsonia 20: 185-194. PERRING, F. H. & FARRELL, L., comp. (1977). British Red Data Books: 1. Vascular Plants. Society for the Promotion of Nature Conservation, Lincoln. PERRING, F. H. & FARRELL, L., comp. (1983). British Red Data Books: 1. Vascular Plants, 2nd ed. Royal Society for Nature Conservation, Lincoln. PERRING, F. H. & GARDINER, B. G., eds (1976). The biology of bracken. Botanical Journal of the Linnean Society 73: 1-302. PERRING, F. H. & RANDALL, R. E. (1972). An annotated Flora of the Monach Isles National Nature Reserve, Outer Hebrides. Transactions of the Botanical Society of Edinburgh 41: 431-444. PERRING, F. H. & SELL, P. D. (1959). Further notes on the flora of Cambridgeshire. Proceedings of the Botanical Society of the British Isles 3: 165-171. PERRING, F. H. & SELL, P. D., eds (1968). Critical supplement to the Atlas of the British Flora. Thomas Nelson & Sons, London. PERRING, F. H. & WALTERS, S. M., eds (1962). Atlas of the British Flora. Thomas Nelson & Sons, London. PERRING, F. H. & WALTERS, S. M. (1971). Conserving rare plants in Britain. Nature 229: 375-377. PERRING, F. [H.] & WALTERS, [S.] M. (1989). The Macmillan Field Guide to British Wildflowers. Macmillan Press, London and Basingstoke. PERRING, F. H., SELL, P. D. & WALTERS, S. M. (1955). Notes on the Flora of Cambridgeshire (v.c. 29). Proceedings of the Botanical Society of the British Isles 1: 471-481. PERRING, F. H., SELL, P. D., WALTERS, S. M. & WHITEHOUSE, H. L. K. (1964). A Flora of Cambridgeshire. Cambridge University Press, Cambridge. SELL, P. [D.] (2004). Obituary: Franklyn Hugh Perring (1927-2003). Nature in Cambridgeshire No. 46: 95- OT: SELL, P. [D.] & MURRELL , [J.] G. (2006). Flora of Great Britain and Ireland. Volume 4. Campanulaceae- Asteraceae. Cambridge University Press, Cambridge. SINKER, C. A., PACKHAM, J. R., TRUEMAN, I. C., OSWALD, P. H., PERRING, F. H. & PRESTWOOD, W. V. (1985). Ecological Flora of the Shropshire region. Shropshire Trust for Nature Conservation, Shrewsbury. SOPER, J. H. & PERRING, F. H. (1967). Data processing in the herbarium and museum. Taxon 16: 13-19. WALTERS, S. M. (1954). The Distribution Maps Scheme. Proceedings of the Botanical Society of the British Isles 1: 121-130. WALTERS, S. M. & PERRING, F. H. (1956). B.S.B.I. Distribution Maps Scheme; Survey of 1955 recording, and suggestions for recording in 1956. Proceedings of the Botanical Society of the British Isles 2: 10-17. WHILD, S. & LOCKTON, A., eds (2005). New Flora of Attingham Park. University of Birmingham & Shropshire Botanical Society, Shrewsbury. WYNNE, G. (1993). Flora of Flintshire. Gee & Son, Denbigh. C. D. PRESTON & P. H. OSWALD 208 OBITUARIES JOYCE LAMBERT 1916-2005 Dr. Joyce Lambert, the first to demonstrate that virtually all the c. 50 lakes in Broadland are man-made, died on May 4 2005. Born in 1916 at Herne Hill, the only child of an estate-agent father, Joyce was educated at the Norwich High School for Girls. Afterwards, she studied botany at the University College of Wales, Aberystwyth, graduating in 1939. Three years later, after a spell as a school teacher in Norwich, she was appointed a lecturer in botany at London University’s Westfield College, now incorporated in Queen Mary College. Her Aberystwyth training had shown her the value of seeing plants in the field and she encouraged her students to get into the field. She had a good eye for spotting, and could communicate the “jizz” of plants stressing the use of vegetative characters, an aspect she put to use in her later ecological work. Prompted by suggestions made towards the end of the Second World War by local naturalist, Ted Ellis, and another Norfolk man, A. R. Clapham, then of Oxford University, she commenced an ecological study of the fens bordering the River Yare in the Surlingham- Rockland area. She published her findings in a series of papers from 1946 onwards, and in 1948 moved to Cambridge University where she received encouragement from _ such luminaries as Harry Godwin and Alfred Steers. While at Cambridge she turned her attention to the fens located in the valley of the Bure, working in partnership with J. N. Jennings, a stratigrapher then based at Leicester University. In 1951 the two of them published in the Journal of Ecology three classic papers relating the alluvial stratigraphy of the Bure valley to the types of vegetational succession which occur there. Jennings meantime had been working on the stratigraphy of other parts of the region, and when writing up his results in the form of a memoir for the Royal Geographical Society (published in 1952 under the title “The Origin of the Broads’) had concluded that most of the broads in the region, if not all, had been formed by natural processes. Meantime Joyce had been carrying out further investigations into the ecology and Stratigraphy of the Yare valley broads and fens. Using a small peat borer she put down a series of closely-spaced cores around various broads in 1952. These revealed to her amazement that- what had previously been thought to be natural lakes had near-perpendicular walls; moreover their floors, some 3 metres or so below the present fen surface, were almost flat. Clearly, these water bodies had originated as peat diggings whose angular shape had been concealed by the overgrowth of fen vegetation once they had become permanently water- filled. Joyce had not made this discovery when she gave her Presidential address to the Norfolk & Norwich Naturalists’ Society in April 1952, but she was able to incorporate her new findings when this was published in that society’s transactions the following year. This startling information, later published in a brief paper in the March 1953 issue of the Geographical Journal, caused a sensation. Indeed, her totally unexpected discovery was sceptically received by those understandably puzzled as to how such extensive excavations could have been dug by hand within areas of the flood plain now subject to regular inundation. To answer these and many other questions, a multi- disciplinary team was established which included a historical geographer, archaeologist and a civil engineer. The joint results of their investigations were published in 1960 in Memoir no.- 3 of the Royal Geographical Society - ‘The Making of the Broads’. Briefly, the team found documentary evidence that proved that substantial amounts of peat for fuel had been dug between the 12th and 14th centuries in a region which was then one of the most economically successful and populous parts of the country. The team did, however, demonstrate that many of the parishes in the region had ‘rights of turbary’ to dig peat within their areas, and that in many cases the boundaries of two, three or more neighbouring parishes are aligned in such a way as to indicate that each of them had rights to extract peat from what later became a single broad. They also concluded that those involved in the industry had considered it worthwhile to excavate to a depth of two to three metres to gain access to brushwood peat, the calorific value of which is superior to that of the material obtainable from near the surface, but that virtually all the excavations in the region had been abandoned by the end of the 14th century as a result of their increasing susceptibility to flooding. OBITUARIES 209 Joyce was appointed a lecturer in botany at Southampton University in 1950, but continued to work closely with the other authors of the R.G.S. memoir. Despite this, she carried out a number of ecological studies on the saltmarshes bordering Southampton Water and other sites, being particularly interested in the spread of the invasive grass, Spartina . Later, she studied the plant communities of the New Forest, dev- eloping in conjunction with Professor Bill Williams a computer-based methodology for their classification. Joyce’s colleagues at Southampton greatly admired her razor-sharp intellect as well as the competence with which she carried out her administrative duties, and she was largely instrumental in setting up a new combined honours degree course in botany and geography. She was regarded by all who knew her as a very caring person. As a consequence of these attributes, she was greatly respected and liked by her students. She retired in 1979 and returned to Norfolk to live at Brundall. She never married. Sadly, Joyce became so physically incapac- itated that she was obliged to spend the last three years of her life in a nursing home at Colney. Nevertheless, she retained her close interest in Broadland, a region now universally regarded as the best example of a lowland wetland system in Britain. One of its particular features of interest is the relationship between its ecology and its land use history, and Joyce’s contribution towards an improved _ under- standing of this will always be remembered. MARTIN GEORGE & CLIVE JERMY RICHARD FITTER 1913-2005 Hands up all of those who learned their flowers from the books of Richard Fitter - or R.S.R. Fitter as he usually signed himself. The words Fitter and field guides surely go together like peaches and cream. Nearly half a century separates his last botanical field guide, Wild Flowers of Britain and Ireland (2003), written with his son Alastair, now Professor of Biology at the University of York, from his first, the famous Collins Pocket Guide to Wild Flowers (1956), written with the late David McClintock. In between he wrote some thirty books, in- cluding a round dozen field guides on plants, birds, pond life and the countryside. Richard Fitter was that now three-star rarity, an all-round naturalist. He knew his birds as well as his flowers, was pretty familiar with insects and other invertebrates, knew London and the Home Counties inside out, and apparently never forgot anything. He once showed me his card-index system, box piled on box, which he used as a reference system. My impression was that he barely needed it. I have a vivid memory of a half-day in the field with him on his home turf near Chinnor in the Chilterns about twenty years ago. He was the only botanist ve known who could clearly identify every chirp and whistle from the bushes while searching for budding Gentianella ciliata with his nose glued to the turf. He knew the history of the local Black Poplars, why they grew where they did, and why they were that shape. He proved incredibly knowledgeable about some piddling plant I was interested in at the time. He seemed, like Linnaeus, a ‘compleat naturalist’. Despite that he was a modest man, without airs and graces. He was the opposite of that contemporary horror, the TV celebrity-naturalist, yet he was a perfectly good lecturer and could spin a good story like the best of them. Merely to list Fitter’s major publications would take up a fair space. His first book was London’s Natural History (1945), one of the first volumes in the famous Collins New Naturalist library. He told me he wrote most of it in the evening after a day at the operational research section of Coastal Command during the war. Instead of simply writing about wildlife localities in the capital he took a novel, thematic approach, examining the influences of trade, smoke, sport and water supply on the city’s wild plants and animals, as well as the role of the blitz in creating a temporarily weed- rich city. Fitter’s relations with Collins blossomed after the war with his authorship of a series of pocket guides, effectively the first modern field guides published in Britain and much influenced by the American bird guides of Roger Peterson. The Pocket Guide to Wild Flowers was the first pocket-sized flora to include nearly all the British wild flowers (apart from grasses and sedges). To help the 210 beginner find their way through the jungle of plants Fitter and McClintock took the bold and controversial step of ranking the flowers by their colour. Fitter had already used a similar approach in his Pocket Guide to British Birds published four years earlier in which birds were grouped by colour, size and habitat. Rarities were marked by stars and the text pruned to the bare essentials of identification. The weak point of Wild Flowers was the illustrations which had to be corralled into plates rather than set next to the text. But despite its limitations The Pocket Guide served a whole generation of British botanists. It was one of those books people fall in love with. Later Fitter wrote a companion volume, Finding Wild Flowers (1972), a guide based on habitats and counties. Richard Fitter was born and bred in London. He was a keen naturalist from boyhood and was particularly interested in birds. As he grew up he became a very active member of the London Natural History Society, surveying starling roosts and heronries in and around London, as well as studying two new species that had begun to nest there, the Black Redstart and the Little Ringed Plover. However, encouraged by his father to take up business studies, Fitter enrolled at the London School of Economics and after graduation became a social scientist, working at the Institute for Political and Economic Planning (PEP) and later Mass Observation. His knowledge of the different worlds of planning and of natural history made him an asset on the Wildlife Special Committee, chaired by Julian Huxley after the war, on which Fitter served as secretary. One of his ‘jobs’ was to visit places the Committee thought worthy of protection to find out what state they were in after six years of war. Even amongst luminaries like James Fisher and Max Nicholson, Fitter’s ability to see wild places in the round was valuable. With his death, the last survivor of that august company of conservation pioneers has passed away. Meanwhile, having married Maisie, his lifelong collaborator and companion, and with two sons and a daughter to feed, he needed more regular work. Fitter became a journalist, helping to edit The Countryman, and for a while acted as ‘open air correspondent’ of The Observer. At the same time the Fitters became a kind of family book business. Among their productions were the popular Penguin Dictionary of Natural History (1967), which Richard wrote with Maisie, and the later Collins Guide to the Grasses, Sedges, Rushes OBITUARIES and Ferns (1984) and Guide to the Countryside (1984) and Guide to the Countryside in Winter (1988), which he did with his son Alastair. He and Alastair also teamed up with Marjorie Blamey to produce the well-known Wild Flowers of Britain and Northern Europe (1974), this time with much better pictures placed opposite to the text. Finally, a generation on in 2003, Richard now aged 91, with Blamey, aged 86, and Alastair produced Wild Flowers of Britain and Ireland which thankfully jettisoned the continental plants and reverted in part to Fitter’s earlier scheme of grouping plants by resemblance and common habitat. Altogether the three “Fitter Floras” offer an evolving record of botanical illustration and identification that is hard to match anywhere. When not writing books, Richard Fitter was a very active conservationist. For a long time he © and Maisie practically ran the Fauna and Flora Preservation Society (now F.F.I., Fauna and Flora International) in Britain. He was also a member of the I.U.C.N. Species Survival Commission, and did a stint as the Council for Nature’s information officer as well as becoming successively secretary and treasurer of the British Trust for Ornithology. He was a founder member of ‘BBONT’, the Bucks, Berks and Oxon Wildlife Trust and was involved with such bodies as the British Deer Society, the Worldwide Fund for Nature and, latterly, the Galapagos Conservation Trust, where first his son Julian, and now his grandson Daniel, worked as a wildlife guide. Yet what may prove to be one of the most important acts of his long career was something Richard Fitter did instinctively and without a thought that it would one day form a valuable record: he kept detailed notebook records. Made over fifty years, his first flowering dates of wild flowers provide an unusual continuous run of data revealing the effects of climate warming. He and Alastair wrote an often quoted paper on it for the journal Science. And shortly before his death at the age of 92 Richard appeared on BBC2s Springwatch programme to talk about it, one of his relatively few television appearances. Richard Fitter was a member of the B.S.B.I. since 1952 and was later made an honorary member. Particularly later in his life he often came to field meetings and conferences. In 1978 he was admitted to the Order of the Golden Ark for his work on threatened species. He also received the Peter Scott medal from the British Naturalists’ Association and _ the OBITUARIES 211 Christopher Cadbury medal from the R.S.N.C. (now the Wildlife Trusts). Never one to be inactive, he was working on three books, including a flora of France, when he died in September 2005. The name of Richard Fitter will long be remembered by the generations of B.S.B.I. members who grew up with McClintock and Fitter, and later turned to Fitter, Fitter and Blamey to sort out problem plants. Few have ever used their natural history skills to better effect, whether to produce books tailored to our needs, to promote good causes or simply to put on public record one’s observations and thoughts on wildlife. There are not many left like him. PETER MARREN ee at, i@ : +e ree : apogee nash a te eas pone > oo litle — ir ta coal | earned gore iin ron _ ans cece * — CI SEN eg 2 ee AHH WW PWNrRN Oomnmmoonordndrinranaran~ ARWNFDOCAMANIDMAKRWH lolly H2. HS) H4. HS. H6. lalife H8. H9. H10. Jalil. ERD: H13. H14. NOR RR RR RS RS West Cornwall Scilly East Cornwall South Devon North Devon South Somerset North Somerset North Wiltshire South Wiltshire Dorset Isle of Wight . South Hampshire . North Hampshire . West Sussex . East Sussex East Kent . West Kent . surrey .. South Essex . North Essex Hertfordshire . Monmouthshire . Glamorgan . Brecknockshire . Radnorshire . Carmarthenshire Dumfriesshire . Kaircudbrightshire Wigtownshire . Ayrshire . Renfrewshire . Lanarkshire . Peeblesshire . Selkirkshire . Roxburghshire . Berwickshire . East Lothian . Midlothian West Lothian . Fife &Kinross South Kerry North Kerry West Cork Mid Cork East Cork Co. Waterford South Tipperary Co. Limerick Co. Clare North Tipperary Co. Kilkenny Co. Wexford Co. Carlow Laois NAMES OF VICE-COUNTIES IN WATSONIA ke Dip Jipy. 24. Ds 26. Dale 28. 7S), 30. Sil 32: 33% 34. 36. Sie 38. 39} 40. Sh 45. 46. 47. 48. 49. 86. 87. 88. 89. 90. Sr 92. 93: 94. 95: 96. 96b. OTF 98. Hus: H16. H17. H18. H19. H20. H21. 22: H23; H24. 25: H26. Hie H28. S (113) Channel Islands ENGLAND Middlesex 54. Berkshire 5) Oxfordshire S10 Buckinghamshire 56. East Suffolk Syl. West Suffolk 58. East Norfolk 59. West Norfolk 60. Cambridgeshire 61. Bedfordshire 62. Huntingdonshire 63. Northamptonshire 64. East Gloucestershire 65. West Gloucestershire 66. Herefordshire 67. Worcestershire 68. Warwickshire 69. Staffordshire 69b. Shropshire 70. South Lincolnshire TA, WALES Pembrokeshire 50. Cardiganshire al Montgomeryshire a2: Merioneth Caernarfonshire SCOTLAND Stirlingshire 99) West Perthshire 100. Mid Perthshire 101. East Perthshire 102. Angus 103. Kincardineshire 104. South Aberdeenshire aamaeeli(() 5) North Aberdeenshire 106. Banffshire LOW Moray 108. Easterness 109. Nairnshire 110. Westerness IEIL Main Argyll 2. IRELAND South-east Galway H29. West Galway H30. North-east Galway H31. Offaly Is yy, Co. Kildare 33; Co. Wicklow H34. Co. Dublin HSS) Meath H36. Westmeath IsBY/ Co. Longford H38. Co. Roscommon 139: East Mayo . H40. West Mayo Co. Sligo North Lincolnshire Leicestershire Rutland Nottinghamshire Derbyshire Cheshire South Lancashire West Lancashire South-east Yorkshire North-east Yorkshire South-west Yorkshire Mid-west Yorkshire North-west Yorkshire County Durham South Northumberland North Northumberland Westmorland Furness Cumberland Isle of Man Denbighshire Flintshire Anglesey Dunbarton Clyde Isles Kintyre South Ebudes Mid Ebudes North Ebudes Wester Ross Easter Ross East Sutherland West Sutherland Caithness Outer Hebrides Orkney Shetland Co. Leitrim Co. Cavan Co. Louth Co. Monaghan Fermanagh East Donegal West Donegal Tyrone Co. Armagh Co. Down Co. Antrim Co. Londonderry =) NOTES INSTRUCTIONS TO CONTRIBUTORS Scope: Authors are invited to submit Papers and Notes concerning British and Irish vascular plants, their taxonomy, biosystematics, ecology, distribution and conservation, as well as topics of a more general or historical nature. Authors should consult the Hon. Receiving Editor for advice on suitability or any other matter relating to submission of manuscripts. Papers and Notes must be submitted in duplicate, typewritten on one side of the paper, with wide margins and double-spaced throughout. Pages should be numbered. Submission of final edited copy on computer disc will be requested, but two hard copies of the text are acceptable if computer facilities are not available. Format should follow that used in recent issues of Watsonia. Underline where italics are required. Names of periodicals should be given in full, and herbaria abbreviated as in British and Irish herbaria (Kent & Allen 1984). The Latin names and English names of plants should follow the New Flora of the British Isles (Stace 1997). Further details on format can be obtained from the Hon. Receiving Editor or by viewing the website at: http://www.bsbi.org.uk/new_style_manual.htm Tables, figure legends & appendices should be typed on separate sheets and attached at the end of the typescript. Figures should be drawn in black ink or be laser-printed and identified in pencil on the back with their number and the author’s name. They should be no more than three times final size, bearing in mind they will normally be reduced to occupy the full width of a page. Scale-bars are essential on plant illustrations and maps. Lettering should be of high quality and may be done in pencil and left to the printer. Black and white photographs can be accepted if they assist in the understanding of the article. If you are able to submit figures on disc please contact the Receiving Editor to check they are in a suitable format. Contributors must sign a copyright declaration prior to publication which assigns the copyright of their material to the Botanical Society of the British Isles. Twenty-five offprints are given free to authors of Papers and Notes; further copies may be purchased in multiples of 25 at the current price. The Society takes no responsibility for the views expressed by authors of Papers, Notes, Book Reviews or Obituaries. Submission of manuscripts Papers and Notes: Mr M. N. Sanford, c/o The Museum, High Street, Ipswich, Suffolk, IP] 3QH. Books for Review: Dr C. D. Preston, Biological Records Centre, Monks Wood, Abbots Ripton, Huntingdon, PEI7 2LS. Plant Records: the appropriate vice-county recorder, who should then send them to Mike Porter, 5 West Avenue, Wigton, Cumbria CA7 9LG. Obituaries: Mrs M. Briggs, 9 Arun Prospect, Pulborough, West Sussex, RH20 IAL. Back issues of Watsonia are available from the official agents for BSBI Publications, Summerfield Books (John & Sue Atkins), Summerfield House, High Street, Brough, Kirkby Stephen, Cumbria CA17 4AX. Tel: 017683 41577. Fax: 017683 41687. E-mail: bsbipubs @ beeb.net Watsonia August 2006 Volume twenty six Part two Contents LANSDOWN, R. V. Notes on the water-starworts (Callitriche) recorded in Europe ... SQUIRRELL, J. HOLINGSWORTH, P. M., SEARS, J., BANKS, B., FERRY, B. & TH DE GRAAF, D. Assessment of ae es in populations of Coe — L: (Asteraceae) : SHIMWELL, D. W. A Shoddy Tale: perspectives on ‘ eel Gi lara x Wes Yorkshire in the twenty-first century RICH, T. C. G. Conservation of Britain’s biodiversity Hieracium | riddelsdelli (Asteraceae), Riddelsdells’s Hawkweed Es LANSDOWN, R. V., STEWART, N. F., KITCHEN, C. & KITCHEN, M. A. R. The status and conservation of stoneworts in West Gloucestershire (v.c. 34) and North Somerset (v.c. 6) FOLEY, M. J. Y. J. H. Penson: mystery man and mystery eon PLANT RECORDS OBITUARIES Botanical Society of the British N Isles ISSN 0043-1532 Typeset by D. K. & M. N. SANFORD Printed in Great Britain by PALLADIAN PRESS, UNIT E, CHANDLERS ROW, PORT LANE, COLCHESTER, ESSEX CO1 2HG 105-120 121-126 127-137 139-144 145-169 171-178 179-195 197-211 AILSON!I Journal of the A Botanical Society of the British Isles Volume 26 Part 3 February 2007 Editors: M. Briggs, M.J. Y. Foley, D. L. Kelly, D. R. McKean, M.S. Porter, C. D. Preston, M. N. Sanford Botanical Society of the British sles Registered Charity Number: 212560 Applications for membership should be addressed to the Hon. General Secretary, c/o Department of Botany, The Natural History Museum, Cromwell Road, London, SW7 5BD, from whom copies of the Society’s Prospectus may be obtained. Officers for 2006—2007 President, Dr R. J. Gornall Vice-Presidents, Prof. C. A. Stace, Prof. A. J. Richards, Mr A. O. Chater, Dr C. D. Preston, Dr L. Farrell Honorary General Secretary, Mr D. A. Pearman | Honorary Treasurer, Mr M. E. Braithwaite Editors of Watsonia Papers and Notes, M. J. Y. Foley; D. L. Kelly, D. R. McKean, M. N. Sanford* Book Reviews, C. D. Preston Plant Records, M. S. Porter Obituaries, M. Briggs *Receiving editor, to whom all MSS should be sent (see inside back cover). © 2007 Botanical Society of the British Isles The Society takes no responsibility for the views expressed by authors of Papers, Notes, Book Reviews or Obituaries. ; The cover illustration of Primula scotica Hook. (Scottish Primrose) was drawn by Rosemar Wise. 23 Max Walters Memorial Issue of Watsonia This issue of Watsonia is dedicated to the memory of S. M. Walters (1920-2005). It is fitting that B.S.B.I. should commemorate Max’s immense contribution to British and European Botany in this way. Max was an inspiration to all the many B.S.B.I. Projects in which he was involved. He will probably be best known to British botanists as co-editor with Frank Perring of the landmark AZlas of the British Flora, published in 1962. He belonged to that great generation of phytogeographers that included Max’s close friends Frank Perring and David Webb, who quite literally put British Botany on the map. Max’s interests were, of course, much wider than distribution mapping and the bibliography (pp. 220-227) gives an idea of the broad range of material that he published over some 60 years. Indeed, his interests were so wide-ranging that it was relatively easy to pull together a set of appropriate papers for this memorial issue. His work ranges from the parochial to the international; from single species studies to overviews of plant taxonomy; from historical to modern day; all of it written in an elegant, self-effacing style which never allows the author’s ego to get in the way of the information. It is notable how much of his publishing was as a joint author or collaborator - indeed, in his Presidential Address to the B.S.B.I. in 1974 he spoke of the “real pleasure” he gained from collaborative work. This was a reflection both of his generous nature and of a deeply felt belief in co-operation at both the national and international level. The task of putting this issue together was made easier through the generous assistance of members who either suggested relevant work or made contributions themselves. I am grateful to Chris Preston for his suggestion that we produce this memorial issue and for his contribution on species at the northern or southern edges of their European range in the British Isles - a subject of particular relevance given Max’s involvement in Flora Europaea. Chris has also been involved in Mark Gurney’s studies on Oxlip hybridisation in Cambridgeshire; together with Max’s long- standing friend David Briggs, they have shed some light on a subject Max himself had examined in 1968. Joachim Kadereit, one of Max’s PhD students, has kindly contributed a paper on European phylogeography. Peter Sell has put together an excellent obituary and bibliography. Kevin Walker has followed up some of Max’s predictions from 1970 published in The flora of a changing Britain. David Pearman has explained the thinking behind the categorisation of neophytes in the New Atlas; the links with historical botany and garden literature are very relevant to Max’s interests. Several other papers in this issue were already in the pipeline before we had considered dedicating it to Max. Nevertheless, Teucrium scordium in Cambridgeshire, the taxonomy of Eleocharis mamillata and the history and contents of the Hull Herbarium are all of particular relevance to Max’s work. Max’s son, Martin has kindly provided the photograph overleaf, taken by Philip Mynott in June 2000. I am sure Max would be pleased to herald the use of colour in this journal. 2 MARTIN SANFORD R 2 Editor, Watsonia Pd eae non SORES Max Walters in the wildflower garden of Great St Andrews Church, Cambridge, in June 2000. Watsonia 26: 215-227 (2007) UN) Obituary STUART MAX WALTERS (1920-2005) Max Walters was a leading light in the world of plant taxonomy and the conservation of the countryside, and he was renowned for his co- operation with fellow-workers in continental Europe for over fifty years. He had a natural gift for bringing together people from all walks of life for the common cause. I have a vivid memory of him with a group of people so brought together to perform the first coppicing in Hayley Wood for over forty years: most of them had never before used the motley array of saws, axes, Slashers, billhooks, sickles and scythes, but the job was done. Max was born in Oughtibridge near Sheffield on 23 May 1920; his father Bernard was a furnace foreman in a steel rolling mill who had been apprenticed to his own father, and his mother was a teacher. Soon after 1920 they moved a short distance to Stocksbridge, where Max spent his boyhood. Both towns lay in the smoke-filled valley in what was termed ‘the Socialist Republic of South Yorkshire’. The Grammar School that he attended at Penistone, between Sheffield and MHuddersfield, was founded in 1392. He went up to St John’s College, Cambridge, in 1938 with an Open Exhibition from the College and a County Major Scholarship from the West Riding. His elder brother Alec, a mathematician, had preceded him _ to Cambridge at Peterhouse. Max was promoted to an Open Scholarship after one year and gained First Class Honours in Part I of the Natural Sciences Tripos in 1940. From his parents Max inherited the deep Christian faith and socialist principles that guided his whole life. As a pacifist, he spent the War Years from 1940 to 1945 working in hospitals, first in Sheffield and then near Bristol. Although he had been fascinated by plants and their variation since childhood, he left school expecting to become a_ biochemist. However, he was greatly impressed by a trip to the Swiss Alps made between school and university on a travelling scholarship from the West Riding, by a visit to the Cairngorms on a high-powered field trip from the Cambridge Botany School, and especially by the teaching at Cambridge of Humphrey Gilbert Carter, Director of the University Botanic Garden, and Harry Godwin, who later became Professor of Botany and received a knighthood. On returning to Cambridge in 1945, Max achieved a First in Part II Botany in 1946. Later that year he commenced research for a PhD on a DSIR grant. In the introduction to his dissertation, submitted and approved in 1950, he noted that he had been introduced to the taxonomic problems of Eleocharis which he was studying for his PhD by a Norfolk amateur botanist, Richard Libbey, when they were identifying the Cyperaceae of Norfolk fens. His doctoral work included the relatively new field of experimental taxonomy, which involved cultivating plants in uniform conditions. Much of his experimental work was done at Wicken Fen. He received only light guidance from his supervisor Harry Godwin, who recognised Max’s self-motivation. In 1947 Max spent the summer at Uppsala in Sweden where some of the seminal work in experimental taxonomy was carried out; he had learned Swedish in Cambridge from Brita Mortensen, a granddaughter of the dramatist August Strindberg. Latin, French and German he learned at school and the use of German Floras with Gilbert Carter in practicals in which the class named plants with Garcke’s Flora von Mittel- und Nord-Deutschland. Later he learned Russian. His linguistic abilities were to serve him well in his co-operation with many continental botanists. In 1948 he married his childhood sweetheart Lorna Strutt, who was to be his bedrock for a long and happy family life. In 1949 Max was elected to a three-year Research Fellowship at St John’s. Later in the year he was made Curator of the Herbarium at Cambridge, a post which had been vacant since 1931. In the beginning of 1950 I returned to that herbarium from doing my National Service. The whole of the Botany School was being renovated, its museum disbanded and its herbarium moved to the room occupied by the former museum. The old herbarium was a dull and dismal place with brown cupboards, poor lighting and few visitors. The new herbarium 216 was more spacious, with white cupboards and modern lights. Max was to turn this new herbarium into a hive of activity over the next 20 years as more and more money became available. As well as myself, three young lady technicians worked full-time on the general herbarium and Dorothy Soden part-time on the mosses. Professor E. J. H. Corner was to work on tropical botany and fungi. It was a very happy place to work in. Max devoted much time to teaching. He gave second year students lectures on experimental taxonomy, variously entitled ‘Evolution and reproduction’, “Taxonomy and evolution’ and ‘Genetics and experimental taxonomy’. He continued Humphrey Gilbert Carter’s practice of teaching the third year students the families of flowering plants in a global context, as well as using Garcke’s Flora von Mittel- und Nord- Deutschland to teach them elementary German. Max also followed Gilbert Carter in taking students, and anyone else who asked to come along, on afternoon cycle rides to get to know about the local flora. He gave tutorials to first year students in Botany up to.1964 and in the Biology of Organisms from 1966; this involved a whole range of disciplines from morphology and anatomy through physiology to ecology. In the Easter vacation or in the summer Max took students on excursions covering taxonomy and ecology, mostly abroad, to Sweden (1950), Portugal (1951), Scotland (1953), the Austrian Alps (1954), Wales (1955), Ireland (1959), Bavaria (1961), Slovakia (1963), southern France (1965), Slovenia (1967), Majorca (1969 and 1973) and Montenegro (1971). Max supervised a number of PhD students — Eklas Bari (Silene), Christopher Cook (Ranunculus subgenus Batrachium), Shahina Ghazanfar (Silene), Keith Goodway (Galium), Geoffrey Halliday (Arenaria), Joachim Kadereit (Senecio), Alan Leslie (Ranunculus auricomus agg.), Neville Marchant (Veronica), Pierre Morisset (Ononis), David Ockendon (Linum), Honor Prentice (Silene), Gordon Smith (Potentilla) and Fenella Wrigley (Silene). Alchemilla was probably his favourite genus, followed by Aphanes, Montia, Silene, Betula and Viola. In April 1954, the B.S.B.I. appointed Max as part-time Director of a scheme to map the distribution of British and Irish plants; Frank Perring became the ‘Senior Worker’ in the scheme the following October. The Atlas of the British Flora was published in 1962. From the ~ records accumulated in Cambridgeshiré for the Atlas, with those in the Cambridge Natural OBITUARY S. M. WALTERS History Society's card index, A Flora of Cambridgeshire by Max, Franklyn, Harold Whitehouse and myself was published in 1964. During the International Botanical Congress in July 1954, in a brasserie on the banks of the Seine, an informal meeting of Tom Tutin, Roy Clapham, John Gilmour, Alan Burges, David Valentine and Vernon Heywood (all Cam- bridge students) took a decision that a Flora of Europe could and should be written. In January 1955 David Webb was added to the group and in March 1956 Max was invited to join the committee. The Linnean Society of London acted as sponsors and Cambridge University Press offered to publish it. Volume 1 was published in 1964, Volume 2 in 1968, Volume 3 in 1972, Volume 4 in 1976 and Volume 5 in 1980. As well as editing 23 of the families including the Caryophyllaceae and half of the Compositae Max wrote 19 genera, including Alchemilla with B. Pawloski and Senecio with Arthur Chater. Flora Europaea was one of the most important taxonomic works published in that century; it brought to the Cambridge Herbarium visiting botanists from all over the world and they were regularly entertained by Max and his family. The final conference was held at King’s College, Cambridge, between 31 August and 4 September 1977 and brought a total of 131 visitors from 30 countries. Max had become a Fellow of King’s in 1964 and bore the brunt of the organisation, together with Gigi Crompton. It was probably the last time that great group of Cambridge botanical taxonomists who originated as Gilbert Carter students in the 1930s, which included the editors of Flora Europaea, and those who were Max’s students between 1950 and 1970 came together in one place. As well as the Atlas, Flora Europaea and A Flora of Cambridgeshire Max’s curatorship days brought several other major publications. Two New Naturalist books became bibles for many field botanists — Wild Flowers with John Gilmour, then Director of the Botanic Garden, and Mountain Flowers with John Raven. This reminds me of Max’s height: he was well over six feet and had very long legs, which were similar to those of John Raven, both of whom I had to run to keep up with when going down a mountain. Perhaps the most important of his writings, however, was Plant Variation and Evolution, written with David Briggs, which has held its own at universities and institutes for over 30 years. With B. Pawlowski he wrote the 50-species account of Alchemilla for the - Flora of Turkey. OBITUARY S. M. WALTERS Max also had an active role in the Cambridge Natural History Society. He joined in 1938 and was elected Junior Secretary in 1939 and Senior Secretary in 1940. He was Botanical Curator from 1947 to 1961, Botanical Recorder from 1961 to 1974, President from 1973 to 1975 and Vice-President from 1975 to 1994. He retired from Council only recently, after being continuously on it for 53 years. He exhibited at the annual conversazione for many years. Max gave a huge amount of his time to conservation, especially the Cambridgeshire and Isle of Ely Naturalists’ Trust. He was one of a small band of enthusiasts who established first CAMBIENT and then Nature in Camb- ridgeshire, which was sponsored equally at the start by the Natural History Society and the Naturalists’ Trust. From the inauguration of CAMBIENT in 1956 until 1965 he was Secretary for Cambridgeshire, while Tony Vine was Secretary for the Isle of Ely. He was Vice- President from 1965 to 1975 and President from 1975 to 1986. In 1962 CAMBIENT bought its first major reserve, Hayley Wood, which is now world-famous. With Robert Payne, the Honorary Secretary, Max bore the brunt of organising the appeal for the £5000 needed to buy it. In 1975 it gave him great pleasure to write the introduction to Oliver Rackham’s pioneering book Hayley Wood: Its History and Ecology. Max must be credited also with a significant role in getting the Society for the Protection of Nature Reserves, now the Royal Society of Wildlife Trusts, reinvigorated in the late 1950s, although the chief role was played by Ted Smith of Lincolnshire. When a buffet supper was held on 15 November 1996 to mark the fortieth anniversary of CAMBIENT, Ted Smith made a speech in which he paid special tribute to Max’s part in arranging the meeting on 19- 20 June 1957 which led to the ‘Cambridge Declaration’ on the need for an ‘Association of Trusts’. Max had a pivotal role in the running of Nature in Cambridgeshire. There was a very slender issue in 1985 after none at all in 1984. That it continued was very much due to his determination. Knowing of the late William Palmer's enthusiasm for the journal, he successfully argued for the use of half the money raised in his memory to become a ‘float’ to relaunch it. He persuaded Philip Oswald to continue as Editor and invited him and others to form an independent Editorial Board, which he himself chaired. Over the NG Max Walters Al 1965. | years he contributed 15 standard articles, seven book reviews, five obituaries, five notes or reports and two ‘state of the nation’ reports on Nature in Cambridgeshire. Of all the nature reserves he had an interest in, Wicken Fen, belonging to the National Trust, held a special place in his heart. He first visited it in 1939, by bicycle and boat with Humphrey Gilbert Carter, and he did much work there in his PhD days. He was a member of the National Trust’s local Committee from 1949 to 1995, the Botanical Secretary from 1949 to 1976 and 1986 to 1990 and Chairman from 1979 to 1985, and he continued to advise until 2004. When Upper Teesdale came under threat from being flooded for a reservoir, with Harry Godwin he stoutly defended it as a unique site of relict communities of plants and gave evidence to Parliament in an attempt to save it. When they lost their appeal he sent two trips from the Cambridge Herbarium to collect a series of the plants which grew there and a set of the Teesdale Alchemilla species. When Max became the Director of the Botanic Garden in 1973, its future seemed rather uncertain. Undergraduates rarely went there and specimens sent to the Botany School for practicals had greatly declined. Max encouraged people of all kinds and ages to visit the Garden and emphasised its role in education. He insisted on first-rate labelling of 218 all the species. He relaunched the Friends of the Botanic Garden organisation, which had lapsed after its foundation in the 1930s. He encouraged Roland Randall to write the first modern educational leaflets for the Garden, those on native trees and shrubs being illustrated by Rosemary Nicholls. At the time when Max moved to the Garden, Gigi Crompton, whom he first met over conservation of an orchid meadow at Thriplow, had been given a contract by the Nature Conservancy Council to survey rare species in eastern England. He provided her with a base in the Garden and developed the project by gaining extra funds from the N.C.C. to employ a succession of conservation gardeners to study the responses of these rare species in cultivation. Later Gigi was to help him organise many Cambridge events and excursions. Max supervised the construction of a new display bed of rare plants of the region and provided a home for conservation of rare plants of Majorca and St Helena. He was among the first Directors of the great gardens to concern themselves with conservation. - When visiting the Royal Botanic Garden, Edinburgh, in the summer of 1976, Max talked to James Cullen and found that they both had the same idea about producing a flora of garden plants. So was born the European Garden Flora in six volumes, edited by James, Max and 19 others. Max wrote the accounts of 25 genera. Volume 1 was published in 1986, Volume 2 in 1984, Volume 3 in 1989, Volume 4 in 1995, Volume 5 in 1997 and Volume 6 in 2000. Max also developed an interest in the history of botany, which he tied up with his philosophy of plant classification. Like John Gilmour, he believed that dividing plants into genera was mostly artificial and mainly a matter of convenience, and it was very difficult to persuade him to alter concepts and long-estab- lished names. While at the Botanic Garden he wrote The Shaping of Cambridge Botany in 1981, and in 2001 in his retirement, he wrote with Anne Stow, formerly Head of the Scientific Periodicals Library, Darwin’s Mentor: John Stevens Henslow 1796-1861. For the centenary of the official opening of the new building of the Botany School in 2004, Peter Grubb, Anne Stow and he produced /00 Years of Plant Sciences in Cambridge 1904-2004. From 1982 until his death he was an active Editor of the New Naturalist series, for which he wrote Wild and Garden Plants in 1993.:In 1989 he and Frank Perring produced a popular ° OBITUARY S. M. WALTERS identification book, The Macmillan Field Guide to British Wildflowers, with specially commissioned photographs by Andrew Gagg, which was reprinted in 1994. His last published book, with John and Jill White, Trees: a Field Guide, appeared in 2005. His final publication has just appeared and is a substantial essay to accompany the facsimile edition of John Ray’s The Wisdom of God to mark the tercentenary of that author’s death. Max was a keen supporter of my own and Gina Murrell’s Flora of Great Britain and Ireland, christened “FOGBI’ by Frank Perring. He was always much interested in variation and was quick to realise the significance of early- flowering variants introduced to this country: hence his articles in the Grantchester Parish Magazine on Snowdrops and on Sorbus. He also fully understood the taxonomy of apomicts. He wrote the forewords to Volumes 4 and 5 of FOGBI. My last long talk with him was about how we hoped almost all his work and that of his students on the British flora © since 1950 would come together in the five volumes. His last active part in it was to sit in the Cambridge Botanic Garden on a beautiful June morning and select typical specimens from two large beds for nearly all the British Alchemilla species for me to describe. In his last years Max visited the Botanic Garden once a week, and on another day, with his daughter Stella, the Herbarium, where he spent time clearing up all the odds and ends accumulated over the years. He gave loyal support to the Cambridge U3A Botany Group for many years: from 1990 to 2002 he usually gave the opening lecture or took the members for a spring walk along the Backs. Throughout his life Max was so often the calming influence between two antagonistic bodies, particularly on the numerous comm- ittees on which he served. There never seemed to be a lost cause: he would find some way of keeping it going. He never seemed to be in a hurry and his very calmness always gave him a good start in any argument. Considering that many meetings often took place in the evening, sometimes very late, and the number that he attended, it was a wonder he kept going, but I only once saw him completely exhausted. What helped was that, like his predecessor at the Botanic Garden, Humphrey Gilbert Carter, whenever possible he took a short siesta after lunch. On the summer excursions Sunday was always a day off for everybody to do as they pleased. Max always enjoyed a good laugh. OBITUARY S. M. WALTERS I remember standing with him at a village fair on the Slovakian—Romanian border watching our students go round and round on the merry- go-round singing “On top of old Smokey’, and on another occasion we watched a student rush all over a marsh on a very hot day trying to flush a ventriloqual Corncrake so that she could tick it on her list. He could also let annoyance turn into laughter. Once, when I had kept the busload of students waiting for half an hour in Yugoslavia, he greeted me with ‘Where on earth have you been?’ I replied ‘Watching the antics of a pair of Sombre Tits’, and the bus dissolved into laughter. When the laughter died down, he said ‘There’s no answer to that; we’d better move on.’ He was a great fan of the Goons and of Monty Python, and he once laughed so much at Monty Python’s antics that he pulled an intercostal muscle and had to have treatment. As a final note of appreciation of Max’s great kindness and understanding I have to add a vote of personal thanks. He did everything it was possible to do to help me in a lifelong study of the British flora and in a way which was the embodiment of a great University at a time before it became obsessed with money. From an amazing 2000-mile trip round Ireland in 1952 with Tom Tutin, Tige Bécher, David Webb, Donald Pigott and Roy Clapham, through all the student excursions, both at home and aboard, he nearly always took me with him. Max joined the B.S.B.I. in 1944, served on the Council from 1957 to 1962 and 1966 to 1969, was Vice-President from 1962 to 1966 and President from 1973 to 1975, served on the Conservation Committee from 1958 to 1969, the Maps Committee from 1959 to 1965, the Publications Committee from 1958 to 1969, the Records Committee from 1966 to 1969 and the Wild Plant Protection Working Party from 1965 to 1969, and was vice-county recorder for Cambridgeshire from 1949 to 1961. He was referee for Montia from 1951 to 1986, for Potentilla from 1973 to 1979, for Aphanes from 1950 to 2005, for Eleocharis from 1949 to 2005, for Betula from 1983 to 1995, for UNE) Alchemilla from 1949 to 2005 and for Silene from 1962 to 2005. He was awarded his ScD at Cambridge in 1980, the Royal Horticultural Society’s Victoria Medal of Honour in 1984 and the Linnean Medal of Botany in 1995. He became an Honorary Fellow of the Linnean Society and an Honorary Member of the B.S.B.I. At King’s College he served on the Garden Committee. He sat on the R.H:S. Scientific Committee from 1976 to 1994. Max was at various times active in the Fellowship of Reconciliation, the Campaign for Nuclear Disarmament and Christian Aid. He had been to Sarajevo for a three-month visit on a British Council grant, so, after the civil war, he worked hard with his son Martin for the _ restoration of the Botanic Garden and Museum there. A fountain in that Botanic Garden records his enthusiasm and generosity. During Max’s curatorship years the Walters family lived in a flat at 1 Brookside over- looking the Botanic Garden. Later he was the last Director to live in Cory Lodge in the Garden. In his retirement he lived at Grantchester, where he was Churchwarden from 1995 to 2001 and wrote 207 articles for the Grantchester Parish Magazine, some of which were botanical; a full set of this magazine from March 1986 is available in the Cambridgeshire Collection in Cambridge City Library. I believe that, unless one has read these articles, one does not understand the complete man. Max died on 11 December 2005, aged 85. A thanksgiving service for his life was held at 2.30 p.m. on Saturday, 3 June 2006, in Grant- chester Parish Church. His wife Lorna survives him together with their three children, Philip and the twins Martin and Stella. He will be much missed by his many friends. I am greatly indebted to Peter Grubb for much of the information and dates concerned with the University of Cambridge, to Mary Briggs, Gwynn Ellis and David Pearman for information about the B.S.B.I, and to Philip Oswald and Arthur Chater for general help. PETER SELL 220 BIBLIOGRAPHY Publications of Dr Max Walters (1920-2005) 1944 Walters, S.M. Notes on White-flowered Viola odorata L. in the Bristol District. Proc. Bristol Nat. Soc. 27: 41-45. 1946 Walters, S.M. Observations on varieties of Viola odorata L. Bot. Soc. Exch. Cl. Rep. 13: 834-839. 1948 Walters, S.M. Glyceria declinata Bréb., en forbisedd nordisk art. Bot. Not. 1948: 430-440. Walters, S.M. Alchemilla vulgaris agg. in northern England. The Naturalist 1948: 41-43. 1949 Walters, S.M. Alchemilla vulgaris agg. in Britain. Watsonia 1: 6-18. Walters, S.M. Aphanes microcarpa (Boiss. et Reut.) Rothm. in Britain. Watsonia 1: 163-169. Walters, S.M. Biological flora of the British Isles. Eleocharis R.Br. Journ. Ecol. 37: 192-206. 1950 Walters, S.M. On the vegetative morphology of Eleocharis R.Br. New Phytol. 49: 1-7. Walters, S.M. On Arabis turrita L. in the Fellows’ Garden. The Eagle 54 (no. 237): 148-150. 195i Walters, S.M. The study of plant distribution. In Lousley, J.E. (Edit.) The study of the distribution of British plants, pp. 12-23. 1952 Walters, S.M. Alchemilla subcrenata Buser in Britain. Watsonia 2: 277-278. 1953 Walters, S.M. Review of An introduction to historical plant geography by E.V. Wulff with a translation by E. Brissendon. Watsonia 2: 214-216. Walters, S.M. The continental element of our flora. In Lousley, J.E. (Edit.) The changing flora of Britain, pp. 124-129. Pigott, C.D. & Walters, S.M. Is the Box-tree a native of England? (Exhibit) In Lousley, J.E. (Edit.) The changing flora of Britain, pp. 184-187. Walters, S.M. Montia fontana L. Watsonia 3: \-6. Walters, S.M. Eleocharis mamillata Lindb. fil. and allied species. Pep Schweiz. Bot. Ges. 63: 271- 286. 1954 Pigott, C.D. & Walters, S.M. On the interpretation of the discontinuous distributions shown by certain British species of open habitats. Journ. Ecol. 42: 95-116. Walters, S.M. Distribution maps of the British flora. Nature 173: 1079-1080. Walters, S.M. Veronica spicata L. and V. hybrida L. Proc. Bot. Soc. Brit. Isles 1: 89. Walters, S.M. The distribution maps scheme. Proc. Bot. Soc. Brit. Isles 1: 121-130. Walters, S.M. Sur quelques plants recoltées dans le Nord-Ouest de la France. Monde Pl. 1954: 303-304. Gilmour, J.S.L. & Walters, S.M. Wild Flowers. London. 1955 Perring, F.H., Sell, P.D. & Walters, S.M. Notes on the flora of Cambridgeshire (v.c. 29). Proc. Bot. Soc. Brit. Isles 1: 471-481. Walters, S.M. Review of Nordisk K4rlvaxflora 1 by N. Hylander. Watsonia 3: 179-180. Walters, S.M. Welsh recording for the B.S.B.I. distribution maps scheme. Nature in Wales: 1(2): 68-71. 1956 Walters, S.M. & Lambert, C. [A] M. A reconsideration of Silene caelata C. Reid. New Phytol. 55: 133-135. BIBLIOGRAPHY 221 Walters, S.M. Mapping the British flora. Country-side n.s. 17: 410-414. Walters, S.M. Selinum carvifolia (L.) L. in Britain. Proc. Bot. Soc. Brit. Isles 2: 119-122. Walters, S.M. Eriophorum gracile Roth. Proc. Bot. Soc. Brit. Isles 2: 23-24. Walters, S.M. & Perring, F.H. B.S.B.I. Distribution map scheme survey of 1955 recording and suggestion for recording in 1956. Proc. Bot. Soc. Brit. Isles 2: 10-17. Walters, S.M. & Webb, D.A. Calystegia. Proc. Bot. Soc. Brit. Isles 2: 22-23. Raven, J. & Walters, S.M. Mountain Flowers. London. IC SyT Walters, S.M. Distribution maps of plants: an historical survey. In Lousley, J.E. (Edit.) Progress in the study of the British flora. Arbroath. Walters, S.M. Mapping the distribution of plants. New Biology 24: 93-108. Walters, S.M. Babington’s Yew-tree. The Eagle 100: 35-37. 1959 Coombe, D.E., Perring, F.H. & Walters, S.M. Lythrum hyssopifolia L. Proc. Bot. Soc. Brit. Isles 3: 286-288. Walters, S.M. Heleocharis austriaca Hayek and Glyceria declinata Bréb. Two plants new to the flora of Poland. Frag. Flor. Geobot. 5: 239-244. 1960 Walters, S.M. Suggested treatement for Alchemilla in Flora Europaea. Feddes Repert. 63: 127— Sil Walters, SM. Field Meetings 1958, July 27 to August 3 1958. Tralee, Co. Kerry. Proc. Bot. Soc. Brit. Isles. 3: 461-463. 1961 Walters, S.M. The shaping of angiosperm taxonomy. New Phytol. 60: 74-84. Walters, S.M. Wild Juniper on Fleam Dyke. Nature in Cambridgeshire 4: 40-41. Bradshaw, M.E. & Walters, S.M. A Russian Alchemilla in south Scotland. Watsonia 4: 281-282. Walters, S.M. Review of A cytogenic study of the genus Geum by W. Gajewski. Watsonia 4: 256- EST 1962 Walters, S.M. Generic and specific concepts and the European flora. Preslia 34: 207-226. Walters, S.M. Botanical classification. Aslib Proc. 14: 231-233. Walters, S.M. Atlas of the British flora. Country-side 19: 8-9. Perring, F.H. & Walters, S.M. Atlas of the British flora. London & Edinburgh. 1963 Bradshaw, M.E., Sell, P.D. & Walters, S.M. The nomenclature of Alchemilla minor auct. brit. Watsonia 5: 259-261. Walters, S.M. Botanical nomenclature, punched cards, and machines — a comment. Taxon 12: 249-250. Walters, S.M. Abstracting and indexing data. Webbia 18: 19-22. Walters, S.M. Methods of classical plant taxonomy. In T. Swain (Edit.) Chemical Plant Taxonomy, pp. 1-15. London. Gilmour, J.S.L. & Walters, S.M. Philosophy and classification. In Turrill, W.B. (Edit.) Vistas in Botany 4: 1-22. London. Walters, S.M. Eleocharis austriaca Hayek, a species new to the British Isles. Watsonia 5: 329- 3D Perring, F.H., Sell, P.D., Walters, S.M. & Whitehouse, H.L.K. Exhibits which illustrate some past, present and future floras. Flora of Cambridgeshire. In P.J. Wanstall (Edit.) Local Floras, pp.. 113-114. London. 1964 Chater, A.O. & Walters, S.M. Flora Europaea Notulae. Silene L. Feddes Repert. 69: 46-47. Walters, S.M. et al. (Edits). Flora Europaea 1. Cambridge. Wrote: Betula, Boussingaultia, Corrigiola, Cucubalus, Draba, Erophila, Holosteum, Montia, Papaver (with A.B. Mowat), Petrocallis, Portulaca, Pteranthus (with P.D. Sell), Silene (with A.O. Chater). 222 BIBLIOGRAPHY Walters, S.M. Obituary of Gordon Levesley Smith (1937-1963). Proc. Bot. Soc. Brit. Isles 5: 417— 418. Walters, S.M. Principles of taxonomy. Development in Industrial Microbiology 5: 169-172. Perring, F.H., Sell, P.D. & Walters, S.M. Flora of Cambridgeshire. Cambridge. 1965 Walters, S.M. Introduction to a general discussion on Phenetic and Phylogenetic Classification, Syst. Assoc. Publ. 6: 157-164. Walters, S.M. ‘Improvement’ versus stability in botanical classification. Taxon 14: 6-10. Walters, S.M. Documentation and abstracting for the European flora. Rev. Roum. Biol. Ser. Bot. 10: 123-124. Walters, S.M. Natural History. In Steers, J.A. (Edit.) The Cambridge Region. 1966 Walters, S.M. The taxonomic implications of apomixis. In Hawkes, J.G. (Edit.) Reproductive biology and taxonomy of flowering plants, pp. 162-168. Bingley, F. & Walters, S.M. Wicken Sedge Fen: a typographical and botanical guide. 1967 Godwin, H. & Walters, S.M. The scientific importance of Upper Teesdale. Proc. Bot. Soc. Brit. Isles 6: 348-351. Walters, S.M. Identification of British Eleocharis. Proc. Bot. Soc. Brit. Isles 6: 384-386. Walters, S.M. & Bosman, V.G. Exhibition meeting 1966. Alchemilla faeronsis (Lange) Buser and A. alpina L. Proc. Bot. Soc. Brit. Isles, 7: 83. Walters, S.M. List of vascular plants of Wicken Fen. Crompton, G. & Walters, S.M. Thriplow meadows grazing experiments 6. Nature in Cambridgeshire 10: 21-23. 1968 Walters, S.M. Betula L. in Britain Proc. Bot. Soc. British Isles 7: 179-180. Sell, P.D. & Walters, S.M. Notes on Slovakian Plants. Acta Univ. Comen. Bot. 14: 13-19. Walters, S.M. et al. (Edits.) Flora Europaea 2. Cambridge. Wrote: Acer, Alchemilla (with B. Pawlowski), Aphanes, Linum (with D.J. Ockendon), Potentilla (with P.W. Ball and B. Pawlowsk1) and Radiola. Walters, S.M. Chromosome numbers of European species of Potentilla L. Grex Aureae Wolf. Acta Univ. Comen. Bot. 14: 7-11. Walters, S.M. Review of Botanical Latin by W.T. Stearn. Watsonia 6: 391. 1969 Ockendon, D.J., Walters, S.M. & Whiffen, T.F. Variation within Centaurea nigra L. Proc. Bot. Soc. Brit. Isles 7: 549-552. Walters, S.M. Review of Mountain Flowers in Colour by A. Huxley. Watsonia 7: 181-182. Walters, S.M. Obituary Mr H. Gilbert Carter, Nature 221 No. 5179, 497-498. Walters, S.M. Cambridgeshire ferns - ecclestiastic and ferroviatic. Nature in Cambridgeshire 12: 22-25. Walters, S.M. Report on index to European taxonomic literature, V. Simposio de Flora Europaea 11-13. Briggs, D. & Walters, S.M. Plant variation and evolution. Cambridge. Gilmour, J.S.L., Walters, S.M., & King, C.J. Humphrey Gilbert Carter: A memorial volume. Cambridge. 1970 Ockendon, D.J. & Walters, S.M. Studies in Potentilla anserina L. Watsonia 8: 135-144. Sell, P.D. & Walters, S.M. Exhibition meeting Report for 1968. Two species new to Cambridgeshire. Watsonia 8: 93. Walters, S.M. Dwarf variants of Alchemilla L. Fragm. Flor. Geobot. 16: 91-98. Walters, S.M. The next twenty-five years In Perring, F.H. (Edit.) The flora of a changing Britain, pp. 136-141. BIBLIOGRAPHY 228, Pankhurst, R.J. & Walters, S.M. Generation of keys by computer. In Cutbill, J.L. (Edit.) Data processing in biology and geology, pp. 189-203. 1971 Perring, F.H. & Walters, S.M. Conserving rare plants in Britain. Nature 229: 375-377. Walters, S.M. Taxonomic and floristic aspects of plant conservation. In Davis, P.H. et al. (Edits.) Plant life in south-west Asia, pp. 293-296. Smith, G.L. Augmented by Bozman, V.G. & Walters, S.M. Studies in Potentilla L. I. Variation in British P.tabernaemontani Aschers and P.crantzii (Cr.) Beck ex Fritsch. New Phytol, 70: 607-618. Walters, S.M. Index to the rare endemic vascular plants of Europe. Boissiera 19: 87-89. Bosman, V.G., Smith, G.L. & Walters, S.M. Studies in Potentilla 3. Variation in British P. tabernacmontani Ascherson and P. crantzii (Cr.) Beck ex Fritsch. New Phytol. 70: 607-618. Walters, S.M. Christians and conservation. Reconciliation Quarterly 23-28. 1972 Walters, S.M., & Webb, D.A. Taxonomic and nomenclatural notes on Veronica L. Bot. Jour. Linn. Soc. 65: 266-269. Walters, S.M. Endemism in the Genus Alchemilla in Europe. In Valentine, D.H. (Edit.) Taxonomy, Phytogeography and Evolution, pp. 301-305. Walters, S.M. & Pawlowski, B. Alchemilla in Davis, P.H. (Edit.) Flora of Turkey 4: 80-104. Edinburgh. Walters, S.M. et al. (Edits.) Flora Europaea 3. Cambridge. Wrote: Veronica (with N.G. Marchant & D.A. Webb). Gilmour, J.S.L. & Walters, S.M. Wild Flowers. Ed. 2. London. Adam, P., Birks, H.J.B. & Walters, S.M. A contribution to the study of the flora and vegetation of . the Budva area, Montenegro. Glas. Repub. Zavoda ZasSt. Priv. Mus. Titograd 4: 41-72. 1973 Birks, H.J. & Walters, S.M. The flora and vegation of Barno Jezero, Durmitor, Montenegro. Glas. Repub. Zavada. ZaSt. Priv. Mus. Titograd 5: 5—23. Walters, S.M. Alchemilla. In Green, P.S. (Edit.) Plants wild and cultivated, 48-50. Walters, S.M. Obituary of Nils Hylander (1904-1970). Watsonia 9: 295-296. Walters, S.M. The role of botanic gardens in conservation. Jour. Roy. Hort. Soc. 98: 311-315. 1974 Walters, S.M. Exhibition Meeting 1973. Veronica spicata in Cambridgeshire. Watsonia 10: 232. Walters, S.M. The rediscovery of Senecio paludosus L. in Britain. Watsonia 10: 49-54. Edmonds, J.M., Sell, P.D. & Walters, S.M. Some British chromosome counts in the Compositae subfamily Chichorioideae. Watsonia 10: 159-161. 1975 Gilmour, J.S.L., Walters, S.M. & King, C.J. Humphrey Gilbert Carter: a memorial volume. Walters, S.M. Obituary of Ian Hepburn. Nature in Cambridgeshire 18: 4—5. Walters, S.M. Traditional methods of biological identification. In Pankhurst, R.J. Biological identification with computers, 3-8. Walters, S.M. Presidential address 1974. Botanical Co-operation. Watsonia 10: 325-330. Walters, S.M. Silene L. 168-170; Betula L. 299-300; Veronica L. 371-372; Tragopogon L. 430- 431. In Stace, C.A. (Edit.) Hybridization and the Flora of the British Isles. London. Walters, S.M. Two famous medicinal herbs in ‘new’ Cambridgeshire localities. Nature in Cambridgeshire 18: 16-18. Walters, S.M. & King, C.J. European floristic and taxonomic studies. 1976 Walters, S.M. The conservation of threatened vascular plants in Europe. Biological Conservation 10: 31-41. Walters, S.M. Review of Det Grénne Gronland by T.W. Bocher. Watsonia 11: 171. Perring, F.H. & Walters, S.M. Atlas of the British flora. Ed. 2. London and Edinburgh. Walters, S.M. et al. (Edits.) Flora Europaea 4. Cambridge. Wrote: Adoxa L. and Senecio L. (with A.O. Chater). 224 BIBLIOGRAPHY 1977 Walters, S.M. A guide to the University Botanic Garden. Walters, S.M. Review of Flora Republicae Popularis Bulgaricae 6. (Edit. by D. Jordanov). Watsonia 11: 417. Walters, S.M. The role of European Botanic Gardens in the conservation of rare and threatened plant species. Gdrtnerisch-Botanischer Brief (Hamburgh) 51: English pp. 2-22, German pp. 22-43, French pp. 43-62. 1978 Walters, S.M. British endemics. In Street H.E. (Edit.) Essays in plant taxonomy, pp. 263-274. Walters, S.M. Editorial on European nature conservation. Naturopa 31: 3. Walters, S.M. Review of Henslow of Hitcham by J. Russell-Gebbett. Watsonia 12: 185-186. 1979 Walters, S.M. The University Botanic Garden. Mag. Cambridge Soc. 4: 42-52. Walters, S.M. Conservation of the European flora: Aldrovanda vesiculosa L., a documented case- history of a threatened species. Syst. Bot. Plant Util. Biosphere. Conserv. 72-82. Walters, S.M. Progress in biological conservation in Cambridge. In Smith, D. (Edit.) Landscape towards 2000, pp. 56-58. Walters, S.M. Epilobium lanceolatum Seb. and Mauri — a plant to look for in your garden. — Watsonia 12: 399. Walters, S.M. Education in the modern botanic garden. The Garden 104: 374-379. Walters, S.M. The eastern England rare plant project in the University Botanic Garden Cambridge. In H. Synge & H. Townsend, Survival or Extinction, pp. 37-46. Donald, D. & Walters, S.M. Some plants from the conservation section of the University Botanic Garden, Cambridge. Watsonia 12: 192. Walters, S.M. The role of Mediterranean botanic gardens in plant conservation. Webbia 34: 109- 116. Donald, D. & Walters, S.M. Exhibition Report: some plants from the conservation section of the University Botanic Garden, Cambridge. Watsonia 12: 392-393. 1980 Walters, S.M. Apomictic endemism in Alchemilla and Hieracium. Watsonia 13: 73. Walters, S.M. Plant refuge in an English garden. Garden 4: 20-23. | Walters, S.M. Apium repens (Jacq.) Reichb.f. Watsonia 13: 170. Walters, S.M. et al. (Edits.) Flora Europaea 5. Cambridge. Wrote: Eleocharis. 1981 Walters, S.M. The future of the countryside. The Cambridge Review 102: 82-84. Walters, S.M. Aims and methods in mapping the endemic vascular plants of the Balkan Peninsula. In S. Kozuharovy (Edit.) Mapping the flora of the Balkan Peninsula, pp. 83—94. Sofia. Walters, S.M. Henslow’s vasculum. Watsonia 13: 369. Walters, S.M. Priorities in rare species conservation for the 1980s. In Synge, H. (Edit.) The biological aspects of rare plant conservation, pp. XXV—XXVili. Walters, S.M. Historical background to the Garden anniversary in University Botanic Garden, Cambridge 150" Anniversary Celebrations 9" to 14 July 1981. Walters, S.M. The shaping of Cambridge botany. Cambridge. Walters, S.M. Review of science and colonial expansion: the role of the British Royal Botanic Gardens by L.H. Brockway. Watsonia 18: 355-356. Briggs, M., Leslie, A.C. & Walters, S.M. Exhibition report: Lemna minuscula Herter, an American duckweed, as a member of the British flora. Watsonia 13: 360-361. 1982 Leslie, A.C., Pannell, C.M. & Walters, S.M. Varieties in Viola odorata L. in Suffolk and Cambridgeshire. Watsonia 14: 73-74. Walters, S.M. Review of Fotoatlas der Alpenblumen by W. Lippert. Watsonia 14: 217-218. Walters, S.M. Some thoughts arising from the publication of a complete catalogue of plants from the Garden. Gdrtn Bot. Briefe 72: 23-26. BIBLIOGRAPHY 225 Rowell, T.A., Walters, S.M. & Harvey, H.J. The rediscovery of the Fen Violet, Viola persicifolia Schreber, at Wicken Fen, Cambridgeshire. Watsonia 14: 183-184. 1983 Walters, S.M. Fen Violets at Wicken Fen, Cambridge, v.c. 29. Watsonia 14: 457. Leslie, A.C. & Walters, S.M. The occurrence of Lemna minuscula Herter in the British Isles. Watsonia 14: 243-248. Perring, F.H. & Walters, S.M. Atlas of the British Flora. Ed. 3. London and Edinburgh. 1984 Walters, S.M. The relation between the British and the European floras. New Phytol. 98: 3-13. Briggs, D. & Walters, S.M. Plant variation and evolution. Ed. 2. Cambridge. Walters, S.M. The role of Botanic Gardens in plant conservation in south-east Asia. Conservation of tropical plant resources 1984: 230-235. Walters, S.M. et al. (Edits.) The European Garden Flora 2. Cambridge. Wrote: Carex, Cyperus, Eleocharis, Lemna, Scirpus, Spirodela, Wolffia and Wolffiella. 1985 Walters, S.M. The utilization of national plant collections. Scientific Horticulture 36: 79-82. Walters, S.M. Lost and gone for ever. Natural World 15: 17-18. Walters, S.M. The problems of wild plant conservation. Publ. Naturhist. Gen. Limburg 35: 56-59. 1986 Walters, S.M. Stellaria palustris — a declining or overlooked species? Ann. Rep. Huntingdonshire Fauna Flora Soc. 38: 8-10. Walters, S.M. The name of the Rose: a review of ideas on the European bias in angiosperm classification. New. Phytol. 104: 527-546. Walters, S.M. The future of Flora Europaea. Veréff. Geobot. Inst. ETH. Stiftung Riibel 87: 91-99. Walters, S.M. Review of God’s Acre: the flowers and animals of the parish churchyard by F. Greenoak. Watsonia 16: 207-208. Walters, S.M. et al. (Edits.). The European Garden Flora. 1. Cambridge. Walters, S.M. Alchemilla: a challenge to biosystematists. Acta Univ. Ups. Symb. Bot. Ups. 27: 193-198. 1987 Walters, S.M. Obituary: John Scott Lennox Gilmour (1906-1986). Linnean 3: 35-36. Akeroyd, J.R. & Walters, S.M. Flora Europaea: the background to the revision of volume one. Bot. Jour. Linn. Soc. 95: 223-226. 1988 Walters, S.M. The Backs of the colleges. Cambridge Review 109: 42-44. Swan, G.A. & Walters, S.M. Alchemilla gracilis Opiz, a species new to the British flora. Watsonia 17: 133-138. Walters, S.M. Note on cultivated Alchemilla species resembling A. mollis. The Garden 113: 535. Akeroyd, J.R. & Walters, S.M. Flora Europaea: Notulae Systematicae. Taxonomic notes on Silene L. Bot. Jour. Linn. Soc. 97: 338-342. Walters, S.M. The purposes of systematic botany. Symb. Bot. Uppsala 28: 13-20. Walters, S.M. A wild and garden Sagina. B.S.B.I. News 49: 52. 1989 Walters, S.M. Experimental and orthodox taxonomic characters and the deme terminology. PI. Syst. Evol. 167: 35-41. Walters, S.M. Alchemilla mollis (Buser) Roth and related species in gardens. B.S.B.I. News 51: | 23-24. Walters, S.M. (Edits.) The European Garden Flora. 3. Cambridge. Wrote: Agrostemma, Alnus (with K. Ashburner), Arenaria, Betula (with K. Ashburner), Cerastium, Cucubalus, Dianthus (with A.P. Hamilton), Gypsophila, Herniaria, Lychnis, Minuartia, Moehringia, Paronychia, Petrocoptis, Pterorphagia, Sagina, Saponaria, Scleranthus, Silene, Vaccaria. 226 BIBLIOGRAPHY Walters, S.M. Obituary of John Scott Lennox Gilmour. Pl. Syst. Evol. 167: 93-95. (Also two early papers on classification, with a foreword by S.M. Walters.) Walters, S.M. Ferns at Wicken Fen. Nature in Cambridgeshire 31: 61. Perring, F.H. & Walters, S.M. The Macmillan field guide to British wildflowers. London. 1990 Walters, S.M. The taxonomy of garden plants. The naming game. The Garden 115: 324-327. Chater, A.O. & Walters, S.M. Flora Europaea: Notulae systematicae ad Floram Europaeam specialities. Notes on Silene vulgaris (Moench) Garcke and S. uniflora Roth. Bot. Jour. Linn. Soc. 103: 213-216. 199] Walters, S.M. Alchemilla update. The Garden 116: 62-63. Walters, S.M. Review of Atlas Florae Europaeae 8: Nymphaeaceae to Ranunculaceae, edited by J. Jalas and J. Suominen. Watsonia 18: 326-337. Walters, S.M. Ray’s Butterbur still in Paradise. Nature in Cambridgeshire 33: 25-26. Walters, S.M. Review: I.P. Schonfelder (Trans. C.J. King) Wild flowers of the Mediterranean. Nature in Cambridgeshire 33: 41. Walters, S.M. King’s College Fellows’ Garden. Cambridge. Walters, S.M. Wall Lettuce, Mycelics muralis in the Cambridge area. Nature in Cambridgeshire. 33: 54-59. Walters, S.M. Professor Maria Skalinska: an appreciation from England. Polish Bot. Stud. 2: 17— 18. 1992 Walters, S.M. An Alchemilla newly discovered in Norway: might it be in Britain. B.S.B.I. News 60: 68. Walters, S.M. Alchemilla in The New Royal Horticultural Society Dictionary of Gardening 1: 100-104. Walters, S.M. W.T. Stearn: The complete naturalist. Bot. Journ. Linn. Soc. 109: 437-442. Walters, S.M. Notable trees and shrubs in King’s College Fellows’ Garden. 1993 Walters, S.M. A brief history of the Backs, Cambridge Univ. Alumni Mag. 10-11 (Easter term 993) Walters, S.M. Wild & garden plants. London. Dickson, J.H., Huber, R. & Walters, S.M. Alchemilla acutiloba Opiz new to Scotland. Bot. Jour. Scotland 46: 499-502. Walters, S.M. Herbaria in the 21° century: why should they survive? Webbia 48: 673-682. Walters, S.M. Stellaria neglecta Weihe in Cambridgeshire. Nature in Cambridgeshire 35: 61-63. Walters, S.M. et al. (Edits.) Flora Europaea. 1. Ed. 2. 1994 Walters, S.M. Obituary: Professor David Webb. The Independent (11 October 1994). Walters, S.M. Classic wildlife sites: Wicken Fen Nature Reserve. British Wildlife 6: 5-13. 1995 Walters, S.M. The taxonomy of European vascular plants; a review of the past half century and the influence of the Flora Europaea project. Biol. Rev. 70: 361-374. Walters, S.M. Four dwarf Caucasian Alchemillas for the rock garden. The New Plantsman 2: 185— 189. Walters, S.M. Obituary: Richard William (Dick) David (1912-1993). Watsonia 20: 320-321. Walters, S.M. et al. The European Garden Flora. 4. Cambridge. Walters, S.M. & Oswald, P. Persicaria species at More Fen, Swavesey. Nature in Cambridgeshire 37: 46-47. 1996 Walters, S.M. et al. Cambient 40th Anniversary. 15 November 1996. Walters, S.M. News from the Sarajevo Botanic Garden. Friends of the Botanic Garden newsletter 41: 19-21. BIBLIOGRAPHY 227 Walters, S.M. Obituary: David Allardice Webb (1912-1994). Watsonia 21: 3-7. Walters, S.M. David Webb’s contribution to the distribution maps scheme. Watsonia 21: 7. Walters, S.M. Swamp Stonecrop, Crassula helmsii in Cambridgeshire. Nature in Cambridgeshire 38: 24-26. Hedberg, O. & Walters, S.M. Optimizing the use of herbarium resources. Taxon 45: 93-94. 1997 Walters, S.M. Botanic records and floristic studies. In Friday, L.E. (Edit.) Wicken Fen the making of a wetland nature reserve. Friday, L.E., Walters, S.M. & Lock, J.M. Carr and woodland. In Friday, L.E. (Edit.). Wicken Fen the making of a wetland nature reserve pp. 82-97. Walters, S.M. & Preston, C. Profiles of new Honorary Members, Peter Sell. B.S.B.I. News 76: 7-8. Walters, S.M. Eleocharis quinqueflora at Wicken Fen. Nature in Cambridgeshire 39: 53-54. Walters, S.M. et al. (Edits.) The European Garden Flora. 5. Cambridge. Wrote Viola. Briggs, D. & Walters, S.M. Plant variation and evolution. Ed.3. 1998 Walters, S.M. Review of Cultivating women, cultivating science, Floras’ daughters and botany in England 1760-1860 by A.B. Shteir. Watsonia 22: 200-201. 1999 Walters, S.M. The changing flora of Cambridgeshire. B.S.B.I. Druce lecture. Cambridge 6 February 1999. B.S.B.I. News 81: 65—70. 2000) = Walters, S.M. U3A excursion to Knapwell Wood. Nature in Cambridgeshire 42: 68. Walters, S.M. et al. (Edits.) The European Garden Flora: 6. Cambridge. Wrote Veronica and Veronicastrum. Walters, S.M. History of the Dahlia. In C. Jennings, Samuel Widnall and His Dahlias. 2001 Walters, S.M. & Stow, E.A. Darwin’s Mentor: John Stevens Henslow, 1796-1861. Cambridge 2004 - Walters, S.M. Grubb, P. & Stow, E.A. 1/00 years of Plant Sciences in Cambridge 1904-2004. Cambridge. 2005 Walters, S.M., White, J. & White, J. Trees: a field guide. Walters, S.M. Essay to accompany the Ray Society’s facsimile (publication 167) of John Ray’s Wisdom of God manifested in the works of the Creation. Special publication of the Ray Society, London. 2006 Cullen, J. & Walters, S.M. Flowering plant families: how many do we need? In Leadley, E. & Jury, S.L. (Edits.) Taxonomy and plant conservation pp.45—90. A BOTANICAL SAMPLE OF THE 207 POPULAR ARTICLES IN THE PARISH MAGAZINE, GRANCHESTER. A fine Snowdrop year! March 1994. Bindweeds and Morning Glory. October 1994. Snowdrops. February 1995. Flowering Cherries and Plums. April 1995. Speedwells. May 1997. Violets and Pansies. May 2001. Dandelions and their relatives. September 2001. The Swedish Whitebeams on Pearce’s Green. November 2004. Watsonia 26: 229-238 (2007) U9. Determinants of phylogeographic structure: a comparative study of seven coastal flowering plant species across their European range J. W. KADEREIT* and E. WESTBERG Institut fiir Spezielle Botanik und Botanischer Garten, Johannes Gutenberg-Universitdt Mainz, D-55099 Mainz, Germany ABSTRACT We here synthesize our own mostly published work on the phylogeographic structure of seven flowering plant species (Cakile maritima, Calystegia sold- anella, Eryngium maritimum, Salsola _ kali, Halimione portulacoides, Triglochin maritima and Crithmum maritimum) widely distributed along the European coasts. Three categories of factors were identified to determine phylogeographic structure and are illustrated with examples. These are 1) historical abiotic factors such as Quaternary climatic changes resulting in sea level changes which may explain, e.g., a phylogeographic break in the Dardanelles and Bosporus straits, 2) modern abiotic factors such as sea currents important for the dispersal of fruits or seeds of most of the species investigated which may explain, e.g., a phylo- geographic break in the Strait of Gibraltar, and 3) factors related to the specific biology of the species which may explain, e.g., the absence of phylo- geographic structure in Calystegia soldanella resulting from clonal growth and _ long-distance dispersal. These factors also act in all possible combinations. The combination of 1) and 2) may be responsible for differential magnitudes in different species of the Dardanelles and Bosporus genetic breaks, that of 2) and 3) for interspecific differences in the Gibraltar break, and 1) and 3) may explain the existence of phylogeographic breaks along the north coast of the west Mediterranean basin present in only some species, and also the deviant behaviour of Triglochin maritima which colonized northern European coasts from inland refugia. KEYWORDS: phylogeography, coastal plant species, glacial distribution areas, sea water dispersal. INTRODUCTION In recent years, the development of appropriate molecular methods has enabled the analysis of the geographical distribution of intraspecific genetic variation. The phylogeographic struct- ure most commonly found is interpreted as the result of range shifts in response to Quaternary *e-mail: kadereit@uni-mainz.de climatic oscillations (reviewed in, e.g., Soltis et al. 1997; Comes & Kadereit 1998, 2003; Taberlet et al. 1998; Hewitt 2000, 2004; Abbott & Brochmann 2003; Brochmann et al. 2003; Stehlik 2003; Tribsch & Schénswetter 2003; Lascoux et al. 2004). Intraspecific genetic structure, however, is also likely to be deter- mined by species-specific biological attributes (Hamrick & Godt 1989; Nybom & Bartish 2000), and by modern barriers to gene flow. These three categories of factors influencing phylogeographic structure are not easy to separate. In this context, we chose to investigate the geographical distribution of intraspecific variation in seven widely and largely co- distributed European coastal plant species of partly contrasting biology. We believe that coastal plant taxa offer several advantages when trying to infer distributional history and recognize the effects of current barriers to gene flow and of species-specific biology. First, they often are azonally distributed and have both latitudinally and longitudinally large geo- graphical ranges. These ranges, more than in narrowly distributed taxa, are likely to still contain both refugial and recolonized areas. Second, coastal species have an essentially linear distribution range. This limits the spatial options for migration and facilitates the reconstruction of distributional limits in Quaternary glacials. The seven plant taxa investigated are Sea Rocket Cakile maritima Scop. (Brassicaceae), an annual often growing in the drift-line of sandy beaches or on shingle, Prickly Saltwort Salsola kali L. subsp. kali (Chenopodiaceae), another annual from essentially the same habitat as Cakile maritima, Sea Holly Eryngium maritimum L. (Apiaceae), a perennial from mostly stabilized sandy habitats such as dunes, Sea Bindweed Calystegia (= Convolvulus) soldanella (L.) R. et Sch. (Convolvulaceae), a J. W. KADEREIT & E. WESTBERG 230 ‘(WINUWALADUL “5 “DULIPLADUL *D) Wwe.ys sn.todsog ay) (9) pure (DUD “D) Wes Sso[fouRpsed dy) (¢) ‘(PY ‘§ ‘Saploovjnjsod “FT “WINWAJIADUL “FY “WNUIPIADUL “AD “DUIPIADUL *D) 99991Q YANOS (Pp) ‘(1]DY “G ‘Saploovjnj4od ‘H Swnunjiou “Ad ‘puajiapu “D) yey yyNos (¢) “(Saploopjnjsod “FT “wnunjldU “AD “DUIADW *D) BOURI IOMYINOS (Z) “(1]DY °S ‘Saploovpjnjsod + ‘wmnwipaDU “q “DUJIADUL “D) TeYRIGIO JO Weg oy) (7) oe sdes osoy) JO UONRdO] OY, ‘(XIS 0) DUO WOJJ posoquINU) soul; yory Aq poyeorpul ore sdes onouss o1y1oodse.qut Jo suonisod oy) 1/Dy “¢ pue saploovjnisod “FT “WNUNPLADUL “AD “DUJIADUL “JOY “(MOOG 99S) DINSIY IY} UL poye1odsoouL UIdq DALY S}[NSo1 oy) pue Ajayeredas pozAyeue Ud9q JABY S}IL.YS VIS WO] WNWILADUL “J JO eyep DHOUdS UOR[Ndod oy,L ‘[OQUIAS OURS OY} YIM POYIeU IIe STeNPIAIPUL JUINNSUOD OY) JO SaNIPeoo] Surjduwies oy) pur sloysnjo yong “vase dATsayoo AT[VoTydesdoos & WOIJ SUTJCUISIIO S[RNPIAIPUL JO S19}SN[D d19U0d JO} po}oodsul sem wieIsOUDYd FY OY], “WNUWIDU “Fy YVIA pottyduioxa SI SISAyeUe OY], “S[eENPIAIPUL UIMJOqG SooURISIP ONOUNS SI] pue ION Jo sisAyeue Sululo¢ JOQYsION Aq poutejgo sidjsnjo Jo UONNQLYsIp yeorydesso9yH “| AANDI4 005 ' ) sebueys Go0';0 ———__—.--* "a, Fh 6S rf PHYLOGEOGRAPHIC STRUCTURE OF SEVEN COASTAL SPECIES 231 perennial creeper from dunes and_ sandy beaches, Rock Samphire Crithmum maritimum L. (Apiaceae), a perennial from rocky shores, and Sea Purslane Halimione portulacoides (L.) Aellen (Chenopodiaceae) and Sea Arrowgrass Triglochin maritima L. (Juncaginaceae), both perennials from salt marshes. We here summarize our published studies on these seven species (Kadereit et al. 2005; Arafeh & Kadereit 2006; Lambracht er al. 2007) and unpublished work on population level analyses of Cakile maritima and Eryngium maritimum across sea straits between the North and Baltic Seas, the Atlantic Ocean and the Mediterranean Sea, and the Aegean, Marmara and Black Seas (Westberg & Kadereit in prep.). Based on _ previous conclusions, we here offer a new synthesis of our results. MATERIAL AND METHODS PLANT MATERIAL Leaf material of the seven species was collected every 100-200 km along the coast from the Turkish coast of the Black Sea to the southeast coast of Sweden on the Baltic Sea and dried in silica gel to preserve DNA. A few collections from outside this range were also included as well as inland material of S. kali and T. maritima. The latter species is rare in the Mediterranean area and is_ therefore represented from only three localities in that region. A few localities were represented by up to seven individuals in each species (Kadereit et al. 2005), and seven additional populations were sampled in C. soldanella from the same general range (Arafeh & Kadereit 2006). In C. maritima and E. maritimum population samples were taken on a more regional scale around the Bosporus and Dardanelles straits, the Strait of Gibraltar and along the coast of west France, as well as in the Baltic Sea and Kattegat in C. maritima. Usually, ten individuals from ten populations were sampled in each region. AFLP PROCEDURE AND DATA ANALYSIS Details of DNA isolation and the AFLP protocol (amplified fragment length polymorphism; Vos et al. 1995) can be found in Kadereit et al. (2005), Arafeh & Kadereit (2006) and Lambracht et al. (2007). To detect geographical structure of genetic variation, cluster analyses were performed on the AFLP data. In most cases individuals were clustered with Neighbour Joiming (NJ; Saitou & Nei 1987) based on Nei and Li’s genetic distances (Nei & Li 1979). In general bootstrap support for clusters was low. Therefore, the resulting phenograms were visually inspected for clusters or groups of genetically similar clusters corresponding to cohesive geograph- ical areas. Such clusters or groups of clusters contained all or nearly all individuals from a continuous area (see Fig. 1 for more details). For the population samples of C. maritima and E. maritimum individuals were clustered with a Bayesian method implemented in BAPS 3.2 (Corander et al. 2003; 2006). With this method clusters are recognized without using inform- ation on the geographical origin of samples. For more details about methods see Kadereit et al. (2005), Arafeh & Kadereit (2006), Lambracht et al. (2007) and Westberg & Kadereit (in prep.). RESULTS CAKILE MARITIMA, CRITHMUM MARITIMUM, ERYNGIUM MARITIMUM, HALIMIONE PORTULACOIDES AND SALSOLA KALI Little geographic substructure was found along the Atlantic and northern European coasts. Only in C. maritima a genetic cluster dividing Baltic Sea material (including Skagerrak and Kattegat) from the rest was detected. Atlantic — Mediterranean subdivision. In four (C. maritima, E. maritimum, H. portulacoides and S. kali) of the five species sampled from southern Iberia a genetic gap was found between Atlantic and Mediterranean material. This gap coincided with the Strait of Gibraltar in C. maritima, H. portulacoides and S. kali, and included one sample slightly east of the Strait’ in EE. maritimum (Fig. 1). ‘The differentiation between these two clusters was especially marked in E. maritimum and S. kali, whereas in C. maritima and H. portulacoides the genetic gap was of smaller magnitude. In Cr. maritimum no genetic gap was found between the Atlantic and Mediterranean material, and instead the Atlantic cluster traverses the Strait of Gibraltar and reaches the Mediterranean coast of northeast Spain. The population level analysis of C. maritima and E. maritimum from this area also showed a distinct gap between Atlantic and Medi- terranean samples, in both species occurring between Tarifa and Algeciras. As in the large scale analysis, the magnitude of differentiation between the two regions was higher in E. maritimum than in C. maritima. 232 J. W. KADEREIT & E. WESTBERG The Mediterranean. In three species the clustering showed a subdivision of material along the south coast of France (Fig. 1). Samples of H. portulacoides from east Spain appear as a distinct cluster and material from France and western Italy forms several closely related clusters. A similar pattern is seen in Cr. maritimum where the Atlantic cluster reaches northeast Spain. In C. maritima two western Mediterranean clusters overlap in northeast Spain and southern France. In E. maritimum and S. kali the samples from the west Mediterranean basin all fall into the same geographical cluster or the same group of clusters. In all species except E. maritimum there is a subdivision of eastern and _ western Mediterranean samples with Sicily as the approximate geographical border. Italian samples of S. kali from the Ionian Sea fall into the western Mediterranean cluster but otherwise samples from the Adriatic and Ionian Seas fall into their own cluster in C. maritima, Cr. maritimum, H. portulacoides and S. kali. With the exception of C. maritima this cluster additionally includes some samples from the western Aegean Sea. East of the Ionian Sea the Aegean and Black Sea samples form additional geographical clusters in all five species. Although no further subdivision of these clusters was found in the large-scale analysis, the population level analysis from the straits between the Black Sea, Marmara Sea and Aegean Sea revealed clear genetic gaps at the Bosporus strait in C. maritima and E. maritimum as well as at the Dardanelles strait in C. maritima. The amount of genetic differentiation in the Bosporus was higher than in the Dardanelles in C. maritima, but in both cases lower than in E. maritimum from the Bosporus. TRIGLOCHIN MARITIMA In the NJ (Fig. 2) and Bayesian analyses two groups were inferred. One Atlantic group contained samples from the Portuguese, Spanish and French coasts and the British west coast, and another group consisted of material from the North Sea, Baltic Sea and Mediterranean coasts as well as Norwegian and Central European inland material. CALYSTEGIA SOLDANELLA The results from C. soldanella contrasted with those from the other species in that no geographical clusters could be identified in the NJ analysis (Fig. 3). Any genetic cluster recognized in the large scale or population level analyses consisted of geographically widely scattered individuals. DISCUSSION Our comparative analysis of phylogeographic patterns in seven flowering plant species of partly different biology, and the consideration of historical and modern abiotic factors allow us to formulate hypotheses about the possible causes of the patterns observed. DARDANELLES AND BOSPORUS: EVIDENCE FOR THE PERSISTENCE OF HISTORICAL GENE FLOW BARRIERS In the population level analysis of C. maritima and E. maritimum, a distinct genetic gap was found in the Bosporus, and also in the. Dardanelles in C. maritima. This gap was not detected either in these two species or in the other species investigated in the large scale analysis of Kadereit et al. (2005), probably as a result of limited sampling of only individual genotypes in this area. Following Kadereit et al. (2005), both species could have grown along the coasts of this area during the last glacial maximum (LGM). During the Quaternary glacials the sea levels of the Black Sea, the Sea of Marmara and the Aegean Sea were repeatedly lower than the sills of the Dardanelles and the Bosporus (Aksu ef al. 2002). This resulted in the isolation of the three seas from each other during such periods. As a consequence, populations in these areas would have been isolated from each other. On the other hand, currents in both the Bosporus and the Dardanelles are unidirectional outflows from the Black Sea to the Marmara Sea and from here to the Aegean Sea. These currents should result in high westward gene flow. The combination of geological history and modern sea currents in conclusion suggest that the genetic differentiation across the Bosporus (and the Dardanelles in C. maritima) is the result of historical isolation during the LGM. STRAIT OF GIBRALTAR: SEA CURRENTS AS A BARRIER TO GENE FLOW For all species growing in the Gibraltar area except Cr. maritimum a more or less distinct genetic gap was found. This gap is most pronounced in E. maritimum and S. kali, and less distinct in H. portulacoides and C. maritima in the large scale analysis (Kadereit 2 PHYLOGEOGRAPHIC STRUCTURE OF SEVEN COASTAL SPECIES ‘DUNIADUL * J, [ISSOJ JO SOS BYVOIpUT F) pure | SIMI] WL, “[eLajeu puevyur uvsdoing [eNUsD pur [e}SvOd ULISOMION pUL ULdURIIOJIPI SUTRIUODS OsTe Jey) dnoIs vag ONTeg/edS YWON & 0} pousIsse sajdures sayor19 pue dnois onuepy ue 0} SuIsuOsoq s[eNprAIpUl jUasoIdas sarenbs “vujiuplu *[ UL SIa\sN{D JO UONGLySIp jeorydeisoas puv sisAjeue SUTULOL IOQUSION ‘7 ANNOY ‘ ' ‘ - eee Se en J. W. KADEREITT & E. WESTBERG 234 “UISIIO [BOTYdeIsOIS 0} SUIPIOSOR Anos JOU OP S[eNpIAIpUy “YZJaUYpjOsS ‘|D Ul S.1d\sn{o JO UONNGLsIp [eo1ydess008 pue siskjeue Surulos ANOGUSION *¢ AUN WY 0001 00s ¥ pee ee i “) , ~ . ‘ . ’ ' Wes a¢ ‘ ' Sia se ¢ . oe y) , cs D - ’ aw s oid ‘ ~ ’ SR os s } . BOOS Pa TS =a . . . oe ENS. os . 2 oe aaa este ontns wwe, ce SS ET ee de hd Ty ah ain POG . = ‘ =. ' pep Che ¢ ‘ - . “see wwe © --* . ~_ ‘ ~~ RASS . - 1 SA COMO OO CHIE Seite’ ~ ' . SSS ‘ . AN ‘ ‘ ‘ . Oe wen ‘ ' > [id . . .~s oe ‘ 1 - s .™ 4 . . ¢ U ips Sis S58 ® 4 Ge, 7+ ene Pte ' 1 ‘> Secs -- ’ se A Sco sods ‘ of vises ’ ‘ at AEN ‘ ‘ Te a Soe ‘ rs oe ‘ ’ RN ‘ G Se ‘ au . 5 aT, . ai : , x OQ bh 0) ‘ ’ . ‘ , ' S ‘ ‘ ‘ ‘ A ' ‘ . 7 ’ ’ ae ‘ ' ‘ > ina 8s aC ‘ \ oe ; ‘ . , A , ‘ , ' ’ x ‘ \ ‘ , . ‘ . , . y sebueyo So0"0 ‘ Z PHYLOGEOGRAPHIC STRUCTURE OF SEVEN COASTAL SPECIES 25) et al. 2005), and distinct and of geographically identical position in the population level analysis of C. maritima and E. maritimum. Except for H. portulacoides, it is the clearest separation in terms of among region genetic differentiation (Kadereit et al. 2005). The Strait of Gibraltar has been open for the last c. Smy (Krijgisman ef al. 1999), and the coasts of the area were likely to have been inhabitable for all species concerned during the LGM (Kadereit er al. 2005). This implies that only a modern barrier to gene flow across the Gibraltar strait can be responsible for the genetic gap found. This gene flow barrier could be in the form of sea currents. The surface currents in the Strait of Gibraltar are influenced by the tidal differences between the Gulf of Cadiz to the west and the Alboran Sea on the east side of the strait (Rey 1983). During high tide surface water is flowing east into the Mediterranean and at low tide the currents are reversed. Possibly the gene flow barrier arises because the changes in sea current direction make it difficult for seeds to pass the strait. This explanation implies that dispersal of the coastal flowering plant species investigated here is largely by sea currents. Similar explanations have been proposed to explain the existence of a genetic gap in the Gibraltar area in several (but not all) marine organisms investigated (Magoulas et al. 1996; Borsa et al. 1997a, b; Pérez-Losada, et al. 1999; Rios et al. 2002; Bargelloni et al. 2003; Nikula & Vain6la 2003; Waters & Roy 2003; Olsen et al. 2004). The exceptional behaviour of Cr. maritimum, where the eastern Spanish material is very clearly part of the larger Atlantic cluster (Kadereit er al. 2005) is perhaps best explained by a recent colonization of the Mediterranean coast of Spain from the Atlantic coast. This does not explain, however, how Cr. maritimum could surmount the Gibraltar barrier postulated here to act in the other species investigated. CALYSTEGIA SOLDANELLA: CLONAL GROWTH AND LONG-DISTANCE DISPERSAL PREVENT THE FORMATION OF PHYLOGEOGRAPHIC PATTERN Among the species investigated, C. soldanella is exceptional in showing essentially no phylogeographic structure. This was found not only in the large scale analysis, but also in a population level study where individuals from geographically widely separated populations did not group according to population (Arafeh & Kadereit 2006). A postglacial arrival of C. soldanella in the study area as explanation for the absence of phylogeographic pattern was ruled out by Arafeh & Kadereit (2006) on the basis of interglacial fossil evidence for the species in the British Isles (Godwin 1975) and on the basis of the observation that the magnitude of intraspecific genetic different- iation in C. soldanella is as high as in the other species investigated. Instead, Arafeh & Kadereit (2006) argued that the exceptional behaviour of C. soldanella is best explained by its biology. The species shows substantial clonal growth, and seeds have been reported to be able to float in sea water for at least 27 months (Arafeh & Kadereit 2006) and to have 90% viability after 1 year of floating (Arafeh & Kadereit 2006) and 30% viability after 18 months of floating (Ridley 1930). The exceptional floating ability results from a very robust seed coat in combination with an air cavity in the seeds. Arafeh & Kadereit (2006) argued that the absence of phylogeographic structure is the combined result of a high frequency of long-distance dispersal resulting from long floating time and long viability of the seeds, and clone longevity, where the age of clones, however, is unknown. High longevity of clones implies that even if establishment of seeds after long-distance dispersal is rare, the chance of their detection is increased by their long persistence. If this interpretation of C. soldanella is correct, it would illustrate that the formation of phylo- geographic pattern through historical or modern abiotic factors can be prevented by a species’ biology. THE DARDANELLES AND BOSPORUS GENETIC BREAKS: THE INTERACTION OF HISTORICAL AND MODERN ABIOTIC FACTORS As discussed above, distinct genetic gaps are found in C. maritima across the Dardanelles and across the Bosporus. In view of ongoing gene flow in this area, these gaps were interpreted as signatures of the closure of these two straits during Quaternary — glacials. Interestingly, differentiation among regions in C. maritima is stronger across the Bosporus than across the Dardanelles. This, in our opinion, reflects the interaction of modern and historical abiotic factors. Whereas the Dardanelles land barrier was broken c. 12 kyr ago (Aksu et al. 2002), the reconnection of the Marmara Sea to the Black Sea happened c. 9-5 kyrs ago (Bahr et al. 2006; Mudie et al. 2004). Accordingly, erosion of the signal left by the glacial closure of the two straits started some 236 2°5 kyrs earlier in the Dardanelles than in the Bosporus, and this is recognizable in the weaker differentiation across the Dardanelles than across the Bosporus. The comparison of these two areas offers the unique opportunity to calculate rates of gene flow through seed dispersal and to predict when the historical signal recognizable will be completely eroded. INTERSPECIFIC DIFFERENCES IN GIBRALTAR: THE MAGNITUDE OF MODERN BARRIERS TO GENE FLOW DEPENDS ON SPECIES BIOLOGY The population-level comparison of C. maritima and E. maritimum across the Strait of Gibraltar revealed the existence of a distinct phylogeographic gap at exactly the same location. Considering the history of the Strait of Gibraltar, which has been continuously open for the last c. 5 my, and the failure to identify a terrestrial barrier to gene flow in the area, modern sea currents were hypothesized to explain the Gibraltar gap found. When comparing the two species investigated, the gap in C. maritima is noticeably smaller than in E. maritimum. This difference can be attributed to the differential dispersability of the two species, where dispersability in terms of floating time in sea water and viability of seeds is higher in C. maritima than in E. maritimum. This finding illustrates that sea currents in the Gibraltar area are not an unsurmountable barrier to gene flow, and, more importantly in the present context, that the strength of modern gene flow barriers is relative and depends on species-specific dispersability. HISTORICAL FACTORS LEAVE SPECIES-SPECIFIC SIGNATURES Among the species investigated, differences in phylogeographic structure exist in the west Mediterranean basin (southwest Italy/Sicily to Gibraltar; Fig. 1). Material from this area falls into one genetic cluster in S. kali, and into several genetically similar clusters in E. maritimum. In C. maritima, H. portulacoides and Cr. maritimum material essentially separates into two groups, of which a western group (e.g., southeast Spain in C. maritima) is more similar to Atlantic material, and an eastern group (e.g., northeast Spain and west Italy in C. maritima) is more similar to Adriatic/east Mediterranean material. The difference between these two species groups (C. mari- tima, Cr. maritimum, H. portulacoides vs. E. maritimum, S. kali) probably can be explained with species-specific responses to Quaternary climatic changes (Kadereit et al. 2005). Wher J. W. KADEREIT & E. WESTBERG considering the geographical position of June/ July/August temperature isotherms now found at the northern distributional limits of the species investigated (C. maritima subsp. maritima: 12°C, S. kali subsp. kali and E. maritimum: 14°C, H. portulacoides and Cr. maritimum: 16°C) during the last glacial maximum (van Andel 2002), and when assuming that these summer isotherms are either directly, as postulated for 4. portulacoides by Chapman (1950), or indirectly responsible for the northern distrib- utional limit of the species investigated, it is likely that E. maritimum and S. kali could have persisted along the entire north coast of the west Mediterranean basin during the LGM, whereas Cr. maritimum and H. portulacoides had to reatreat south. Retreat along the coasts would have meant retreat to the southwest and southeast. The phylogeographic subdivision of | the latter two species along the north coast of the west Mediterranean basin would then be the result of re-colonization from these two directions. No explanation can be offered for C. maritima which, considering its temperature requirements, might have persisted along the entire north coast but still shows a subdivision similar to that of Cr. maritimum and H. portulacoides. The among-species phylo- geographic incongruencies in the _ west Mediterranean basin thus illustrate that responses to historical climatic changes are species-specific and leave a distinct phylo- geographic signal which may eventually dis- appear through ongoing gene flow. Among the species investigated by us, the most striking example for a species-specific response is provided by Triglochin maritima. Today, 7. maritima can grow much further north than the remaining species, and its northern distributional limit is correlated with the 8°C July isotherm. This, the identification of an Atlantic cluster and a cluster containing all material sampled from the North Sea area, the Baltic Sea area, Central European inland localities, the northern Adriatic Sea coast and the Mediterranean coast of southwest France, and the existence of macrofossils of T. maritima near Earith and Upton Warren in southern England (Fig. 3) dated to approximately 40 000 years ago (Bell 1969), led Lambracht et al. (2007) to postulate that modern populations of this species from the North and Baltic Sea areas originated from inland populations rather than from Atlantic Ocean populations as in the other species investigated. PHYLOGEOGRAPHIC STRUCTURE OF SEVEN COASTAL SPECIES 25) CONCLUSIONS The above hypotheses on the possible causes of phylogeographic structure in the species investigated certainly can be taken to even higher levels of complexity. 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Determinants of phylogeographic structure: a comparative analysis of two coastal flowering plant species across three sea straits of different history and along a continuous stretch of coast. (Accepted October 2006) Watsonia 26: 239-251 (2007) 239 Hybridisation between Oxlip Primula elatior (L.) Hill and Primrose P. vulgaris Hudson, and the identification of their variable hybrid P. xdigenea A. Kerner M. GURNEY* Department of Genetics, University of Cambridge, Cambridge CBI 3EH C. D. PRESTON CEH Monks Wood, Abbots Ripton, Huntingdon, Cambs PE28 2LS J. BARRETT Department of Genetics, University of Cambridge, Cambridge CB1 3EH and D. BRIGGS Department of Plant Sciences, University of Cambridge, Cambridge CBI 3EA ABSTRACT Hybrids between Oxlip P. elatior and Primrose P. vulgaris are commonly recorded in mixed pop- ulations of the two species, but there has been debate as to whether or not backcrossing and introgression occur. Allozyme electrophoresis has demonstrated that backcrossed or F,, hybrids are frequent in a mixed population of the species in Cambridgeshire (v.c. 29). Molecular analysis reveals that the hybrid or backcrossed plants are very variable in morphology, and some cannot be distinguished from Oxlips in the field. However, whilst some hybrids are intermediate in character and some are identical, or almost identical, to Oxlips, all were easily distinguished from Primrose. Oxlip is easily separated from the widespread hybrid between Cowslip and Primrose, P. veris x vulgaris. KEYWORDS: Buff Wood, introgression, Primula veris, Primula Xpolyantha, Primulaceae. INTRODUCTION HYBRIDS BETWEEN YELLOW-FLOWERED PRIMULAS IN BRITAIN All three of the British yellow-flowered primulas (Oxlip Primula elatior, Primrose P. vulgaris, and Cowslip P. veris) interbreed to produce hybrids (Valentine 1975). Of the three hybrids, the most widespread is that between P. vulgaris and P. veris, known as False Oxlip Primula xXpolyantha. This plant caused considerable consternation among the botanists of the 19th Century as they debated whether or not the Oxlip was a British plant (Brown 1842; Doubleday 1842; Hill 1842; Leefe 1842; Moxon 1842; Watson 1842; Gibson 1844). The False Oxlip has been recorded as a native from 87 of the 112 Watsonian vice-counties in Britain (Stace et al. 1993), and its distribution matches the range of the less widespread parent, P. veris, fairly closely (Preston et al. 2002). However, the hybrid is rarely present in large numbers (Clifford 1958). The cross between P. veris and P. vulgaris is more fertile when P. veris is the female parent (Darwin 1869), but even so the seeds are less viable than those produced by intraspecific crosses (Valentine 1955). Despite the high viability of seeds from backcrosses to either parent (Valentine 1955), apparent backcross plants or F5s (crosses between hybrids) are rarely reported in the wild (Clifford 1958; Woodell 1965). As one would expect from the limited range of P. elatior, the hybrids that involve this species have a much more restricted dis- tribution than P. xXpolyantha. The hybrid between P. elatior and P. veris, P. media, is rare. It has been recorded only sporadically moun IN@ridn lesser Uermimil — IS), Cambridgeshire (Preston 1993), East and West Suffolk (Simpson 1982), East Norfolk (Miller Christy 1922), and Huntingdonshire (Wells 2003). The P. elatior x P. veris cross very rarely produces viable seed, but when P. veris *Correspondence address: RSPB, The Lodge, Sandy, Beds SG19 2DL e-mail: mark.gurney @rspb.org.uk 240 is the female parent, a tiny fraction of seeds produced may germinate. Valentine (1952) obtained germination rates of 14/4000 for female P. veris x male P. elatior, and 0/2000 for the reciprocal cross with P. elatior as the female. Offspring have been produced by backcrossing P. media to either parent (Valentine 1952), but the only case of backcrossing in the wild that we are aware of is a rather curious- sounding record by Jermyn (1974) of a “hybrid swarm with no P. veris present and only a few roots of P. elatior near a railway with scattered hybrids and abundant P. veris, and a wood with plentiful P. elatior”’. A triple hybrid between all three species, Primula xmurbeckii, has been reported from Suffolk where it seems to have been most often found in colonies of P. elatior x P. vulgaris, but it is extremely rare (Simpson 1982). There is one record from Cambridge- shire, where an apparent cross between native P. elatior and cultivated P. xpolyantha has been reported (Preston 1993). P. xdigenea, the hybrid between P. elatior and P. vulgaris, has a limited distribution, but it seems to be present at all British sites where both parents occur together. It is certainly in all the woods west of Cambridge that contain both P. elatior and P. vulgaris (Meyer & Meyer 1951), and in all those known to us east of that city (which divides the British range of P. elatior into two separate parts). In contrast to the more widespread P. xXpolyantha, P. xdigenea can be present in large quantities where the parent species meet: in Gamlingay Wood (v.c. 29), Peter Walker (unpublished data) counted 3675 Oxlips, 404 Primroses, and 2131 hybrids in 1992. Populations in woods where hybrids are present have been described as hybrid swarms. The character of the P. elatior x P. vulgaris hybrid is very variable. Simpson (1982) noted that “almost every conceivable form can be found between the species. There are some that might be mistaken for Primroses, or even pure Oxlips”. The large numbers of hybrids, and the presence of plants that are almost, but not quite, identical to either parent, suggested that introgression and _ backcrossing may be frequent. This was disputed by Miller Christy (1922), who was “not conscious of ever having seen a plant which appeared to be a second- cross hybrid”, and by morphological investigations by Valentine (1948) and Woodell (1969). However, Valentine (1961) revised this view after a later study, concluding that “At any rate, at present, introgression is certainly occurring.” ; M. GURNEY, C. D. PRESTON, J. BARRETT & D. BRIGGS The variability of the hybrids, and of P. elatior itself, means that it is difficult to use morphological characters to distinguish between first (F;) and later (F,,) generation hybrids and backcrosses, and this probably explains the difference in opinion as to whether populations are introgressed. Using molecular species-specific markers, we have proved that in the well-studied Oxlip population at Buff Wood, Cambridgeshire (v.c. 29), some plants are indeed introgressed or backcrossed (Gurney 2000). Molecular markers also revealed that some plants that were identified as Oxlips using morphological characters were in fact hybrids. With the interest in hybrids from the new hybridisation and the British flora project (Pearman & Preston 2005), we feel it is timely to review the characters used to identify Oxlips and their hybrids, and to demonstrate the ~ variation that can be found in hybrid swarms. Mixed populations of parent species and putative hybrids are not uncommon in the British flora and have traditionally been investigated by morphological or morph- ometric studies. Examples include species of Betula (Brown et al. 1982), Calamagrostis (Crackles 1995), Dactylorhiza (Heslop Harrison 1949), Gentianella (Rich et al. 1997), and Quercus (Kelleher et al. 2003). In some cases cytological evidence has been used to confirm the identity of hybrids in populations where the putative parents differ in chromosome number, such as in the studies of Eleocharis by Lewis & John (1961). Only in recent years has it been possible to combine morphological and molecular studies to investigate the identify of putative hybrids, for example in Potamogeton (Fant et al. 2003), Trichophorum (Hollingsworth & Swan 1999), Salix (Scottish Montane Willow Research Group 2005), and Schoenoplectus (Fay et al. 2003). In this study the identity of putative hybrids has been confirmed by allozyme electrophoresis and these plants have been used in assessing the morphological differences between the taxa. METHODS MORPHOLOGY The populations of P. elatior, P. vulgaris, and P. xdigenea at Buff Wood, Cambridgeshire (v.c. 29), National Grid Reference TL2850, have been the subject of previous studies of hybridisation (Valentine 1947, 1948, 1961; Walters & Ockenden 1968). Plants in this OXLIP HYBRIDS 241 population were compared with those from single-species populations of P. elatior, P. vulgaris, and P. veris (“pure populations”). They were examined and photographed, and characters useful for distinguishing them were noted. From these, drawings were made to show the main characters of each species. Plants of known genotypes at Buff Wood were also drawn and photographed to record the variation in hybrids and backcrossed plants. Specimens of these were deposited in the herbarium of the Department of Plant Sciences, University of Cambridge (CGE). The Buff Wood plants included 26 hybrids of which 11 were non-F,; hybrids. (Backcrosses are produced by hybrids crossing with one of the parent species, and later generation (F,)) hybrids are formed by crosses between hybrids. Both of these could show exactly the same pattern when examined using allozyme electro- phoresis, so we could not distinguish between them. We therefore use the term “non-F, hybrid” to describe backcrosses and _ later generation hybrids (as distinct from first generation F, hybrids).) POLLEN STAINABILITY One flower was collected from plants selected at Foxley Wood, East Norfolk, v.c. 27, and Wayland Wood, West Norfolk, v.c. 28 (P. vulgaris only), and Hayley Wood (P. elatior only), Cambridgeshire, v.c. 29, and Buff Wood (both species), also v.c. 29, and stored in 70% ethanol. The number of plants used varied between sites (Table 2) because we wanted to use the same individuals sampled for allozyme electrophoresis so that we could be certain of the genetic identity of the plants. Unfortun- ately, some of these plants had ali their flowers removed by herbivores before they dehisced, so not all could be sampled for pollen stainability. Eight months after collection, the fertility of each flower was estimated by counting the number of round, stainable pollen grains in acetocarmine. This method was used by Woodell (1969) and Valentine (1948), but neither gave details of the protocol used. We followed the method in Gurr (1953), using a solution of 0-4 g carmine in 55 ml water and 45 ml glacial acetic acid. An anther was removed from the flower and placed on a microscope slide and a drop of acetocarmine was added. A coverslip was then placed on top and tapped to disperse the pollen grains. A sample of 200 grains was counted and the number of unstained, unswollen grains was recorded. Although staining is not necessarily an accurate measure of fertility, this method has been used to distinguish backcrossed plants in previous studies of introgression in Primula (Woodell 1965), and combining the staining results with the electrophoresis should allow an assessment of its effectiveness. RESULTS MORPHOLOGICAL CHARACTERS OF P. ELATIOR, P. VULGARIS, P. VERIS, AND THEIR HYBRIDS The most useful characters for distinguishing between P. elatior and P. vulgaris, based on plants from pure populations, are shown in Figs 1—2 and Table 1. P. elatior has smaller flowers than P. vulgaris, but there is some overlap. More important are the colour of the flowers and the shape of the markings at their centre, scent, and the hairiness of the pedicels and scape. In the mixed population at Buff Wood, some plants combine characters from both species, and molecular analysis shows. that these are a mixture of F; and non-F, hybrids. Three examples of these are shown: an “intermediate” hybrid (Fig. 3); an “Oxlip-like hybrid” (Fig. 4); and a “Primrose-like hybrid” (Fig. 5). Drawings of P. veris and P. xpolyantha are shown for comparison in Figs 6-7. The variation in hybrids confirmed by electrophoresis is illustrated in Plates 1-6, and photographs of the parents from _ pure populations are shown in Plates 7-8. There is no difficulty in distinguishing between the species, between P. xpolyantha and P. elatior, and between P. xpolyantha and P. xdigenea. Hybrids involving P. veris reveal their parentage by a dark orange streak at the base of the petal, by an inflated, more bluntly- toothed calyx, and by shorter hairs. P. vulgaris appears much more distinct than does P. elatior. Many plants at Buff Wood closely resembled P. elatior from pure populations but they showed one or more features suggestive of P. vulgaris, such as slightly longer hairs than in the pure populations, or larger flowers. If they had been found in pure populations, these would probably have been considered to be just outside the extremes of variation within P. elatior and they more or less grade into it. In contrast, all the plants with P. vulgaris characters, such as large, pale flowers, or shaggy hairs, were either obviously 242 M. GURNEY, C. D. PRESTON, J. BARRETT & D. BRIGGS Variation in ring pattern at centre of flowers: left — narrow, sharply defined ring, centre — broad, slightly angled ring, right —broad diffuse ring, Colour of rings varies from dark yellow to orange. FIGURE |. Primula elatior. Calyx rather narrow. Flowers usually facing to one side. Flowers clear yellow, petals often rather narrow. Leaves hairy, often with distinct petiole, usually rather broad and not reddish at base. Scape shortly hairy. Short pedicels. Scale bar = 1 cm. oN! FIGURE 2. Primula vulgaris. Flowers larger than those of P. elatior, with more rounded petals and usually sharper notches. Very pale, almost greenish, yellow flowers with distinct orange star shape at centre, formed by diamond-shaped mark on each petal. Leaves cuneate, with no distinct petiole, often reddish at base. Flowers in a stalk-less umbel. Very rarely on a scape, which is covered in shaggy hairs. Flowers on long pedicels. Pedicel and calyx covered in long shaggy hairs. Scale bar = 1 cm. OXLIP HYBRIDS 243 FIGURE 3. Primula xdigenea (intermediate). Flowers on longer pedicels than in P. elatior, creating more open inflorescence, and not all facing same way. Flowers variable, but usually paler than in P. elatior, and with pentagon or weakly-formed star at centre. Scape, pedicels, and calyx with long shaggy hairs. Leaves broader than in P. vulgaris, but with less distinct petiole than in P. elatior. Scale bar = 1 cm. FIGURE 4. Primula xdigenea (Oxlip-like). Flowers on short pedicels, all facing same way. Flowers pale yellow with clear pentagon mark at centre. Leaves not truncate and not with obvious petiole, but still within range of variation found in P. elatior. Shortly hairy scape. Scale bar = | cm. 244 M. GURNEY, C. D. PRESTON, J. BARRETT & D. BRIGGS NS See SY) FIGURE 5. Primula xdigenea (Primrose-like). Flowers close to P. vulgaris in colour, in stalkess umbel, but with broad diffuse ring at centre. Flowers like P. vulgaris in size and shape. Pedicels with short hairs. Leaves broader than in P. vulgaris. Scale bar = 1 cm FIGURE 6. Primula veris. Broad, inflated calyx with blunt teeth. Flowers with short, round petals. Deep yellow, with sharp dark orange streak at the base of petals. Short pedicels. Leaves glabrous above, very shortly hairy beneath. Leaves sharply truncate. Scape, pedicels, and calyx with very short crisped hairs. Flowers not often facing all same way. Scale bar = 1 cm. OXLIP HYBRIDS 245 FIGURE 7. Primula xpolyantha. Long pedicels, flowers not all facing same way. Flowers larger than in P. veris, more like P. vulgaris in shape. Pale yellow, with orange pentagon at centre, and dark streak on each petal. May rarely have P. veris-like flowers at base. Leaves rounded to truncate, shortly hairy. Scape and pedicels very shortly hairy. Scale bar = | cm. intermediate between the two species or looked exactly like pure P. vulgaris. An example of a plant that looks superficially like P. vulgaris is shown in Fig. 5, but even this is easily dist- inguished from Primrose by its shortly hairy pedicels and very broad leaves. Therefore, the only possible confusion arises with plants that seem to be close to P. elatior, but could be first or later generation hybrids or backcrosses. POLLEN STAINABILITY The results of the pollen staining are presented in Table 2. The P. elatior samples from the mixed population at Buff Wood and the pure population at Hayley Wood both have very high levels of stainability. This seems to be slightly reduced in the hybrids at Buff Wood, but the P. vulgaris from Buff Wood and from the two pure populations also have lower levels of stainability. Pollen stainability can therefore not be used to differentiate between hybrids and P. vulgaris, but there is rarely any difficulty in distinguishing between them using morphological characters anyway. Pollen stainability may be more useful in dist- inguishing between P. elatior and hybrids and backcrosses, but the sample from the pure population at Hayley Wood includes P. elatior plants that have relatively low pollen stainability, so this measure is not conclusive. Unfortunately the leaves sampled from these plants degraded before allozymes could be extracted, so we could not combine pollen staining results with those from allozyme electrophoresis. Our values for stainability are noticeably lower than those of Woodell (1969), and they have higher standard errors. This seems to be M. GURNEY, C. D. PRESTON, J. BARRETT & D. BRIGGS 246 “UOTIODIIP OUILS ay) Sutoey ‘skeMye jou 1nqG ‘uaJQ ‘odevos v& Jo do} je sjoorpod Woys UO SIOMO] "yy90) UINnIpoul §=YyyIM = MOLI “OSOIUILI ul se Asseys JOAU inq ‘SUO] 0} UINIpes| ‘Ayorvod Jaye yy ‘jeuosejuod jsowye pue posure aq Avul SurYy ‘anU90 ye ZulI o3uesJO 0) MOTOA yep peoiq 0} MOLIeU VY ‘dijxQ ul se snoJsounu os jou pure sjoorpod BuO] UO Udo SsIOMO]A ‘odvos ev YUM AqTyens— "Y}99) SUOT 0} UINIPOW YM “MOLE ‘ASseys 1eodde Aewl ‘Suo, 0} UINIpoys/| ‘QSOIUILIg Ul se A[SUOIS se jou yng ‘Adeos uo “SULI pRolg & aavy AU yng ‘a1JUID OY) ye Ie}S ISNJJIp 10 uoseJuod asueio ue YM UO ‘MOTOA oyed 0} ang ‘diyxo ut uey) sored ‘xAyeo oy) UY) Josuoy Aqjensn = ao1pad ‘odeos & IM Ajores AIO A, "YJ99}) SUOT YIM ‘MOLEN ‘Suo] pue Asseys ‘Adevos JSOW]Y ‘sjejod ay} JO oseq ou) ye spuowrip Aq pouloy anuoo oy) ul uosejuod JO Jes osuelio Uy “MOT[OA ‘dijsMO_ Ur UR) Josuoy Ayjensn § s[ao1ped ‘foquin ue ul Jo odeos e Jo do} oy) We SOMO] ‘QSOIUULIg Ul UR) yoo} JoyunTG YIM pue ‘payeyur Apysiys Ajyensy) ‘dijsmog Ur se dso os jou ynq “1104S ‘dijsMo_ Ur ue) asnjjip osow yonut 9q Aveul sty ysnoy) ‘syejod ay) JO aseq oy) Je Years asuvio ue YIM Alpensye) “MOT[OA aind = ayed = Ay peordAy sour yng ‘MoTJOA ojed 0} “SUOTIOIIP JUdIOJJIP Ul suUToRs pue 3urpeoids ‘skemye jou ynq ‘Ajjensn siOMO]y ‘adeos +48 «jo do, je sjaoiped y10Ys UO SJOMOTJ “yoy JUNTA YIM ‘poye|Juy ‘dsiio pue joys “syejod ay) JO aseq oy) je YeonNs a3urio yep MoeU VY JOUVOSOIO [JU] xA[eg ‘advos pur ‘xAyeo ‘sjaoiped uo sirey qUIOS SIOMOLJ UO SSUDLILIY “MOTJOA ang Aqfensn yng ‘afqeuea AOA pfoo aed AyoA 01 9d doop wooly ‘ayqeiea Alo A ‘moy[ak dooq INO[OS JOMO]Y AOIDIA ‘ pauasipx ‘d SLIDE INA “ pyjupdjodx “dq SIDA “d dijxo dijxQ x som ISOM asoIWLig x dijsmog dijsmog SCGIYdAH AHHL UNV YOLLVIA ‘d dYTXO ‘SIMVITNA ‘d ASOUMWIAd ‘SIVAA VINWIAd dITSMOD ONLLVAVdAS AO SUALOVUVHD NIV TT ATE VL OXLIP HYBRIDS 247 e ; a ‘ = 2° ~ i ab @ Plate 1. The petals of this individual are rather Plate 2. This plant is almost indistinguishable narrow and the flowers are rather large. from P. elatior. Note, however, the pentagonal Otherwise this plant is very similar to P. ring. elatior. : > _ . Rs. &. e Plate 3. Another plant that is very close to P. _—_ Plate 4. An obvious hybrid, with large flowers, elatior, but the pedicels are perhaps too long pedicels, a pentagon at the base of the shaggily hairy for that species. petals, and slightly shaggy hairs on the pedicels. Perhaps superficially similar to P. xpolyantha, but that hybrid has a distinct streak at the base of each petal and much shorter hairs. Pe Plates 14. Primula elatior x P. vulgaris at Buff Wood. The allozyme patterns of these plants is incompatible with that of an F;, and they are either backcrosses or later generation hybrids. 248 M. GURNEY, C. D. PRESTON, J. BARRETT & D. BRIGGS ara Co : oS xo Plate 5. Note large flowers with narrow petals Plate 6. An obvious hybrid. The flowers are and star shape at base of petals, rather shaggy large and pale with a star-pentagon at the base, hairs on scape, and red base to leaves. all facing in different directions, and with narrow petals. The scapes are shaggily hairy and the leaves are narrowed into a petiole. Plates 5—6. P. xdigenea from Buff Wood. These plants have allozyme patterns consistent with them being F, hybrids. 2 , Plate 7. Primula elatior. Note the compact Plate 8. Primula vulgaris. Large, cold-toned flowers and secund inflorescence. flowers with a star at the centre. at Plates 7-8. Oxlip Primula elatior and Primrose Primula vulgaris from pure populations. OXLIP HYBRIDS 249 TABLE 2. POLLEN STAINABILITY IN OXLIPS P. ELATIOR (O), PRIMROSES P. VULGARIS (P), AND HYBRIDS (H) Site Pollen stainability (%) n Mean S.E.M 20-60 61—70 71-80 8 1—90 91-95 96-100 Hayley (O) 1 l 1 8 11 96-1 1-89 Buff (O) 1 1 8 10 96:8 LG Buff (H) 1 2 2 1 6 88-8 3:49 Buff (P) 1 4 2 2 9 89-7 2°36 Foxley (P) 1 1 2 1 1 6 V2 86:3 4-74 Wayland (P) 1 1 2 D) 4 DD VW 80-5 6:26 Marley (P) 60 60 99-9 0-01 Lawn (O) 1 1 1 7 60 98-9 0-66 Hayley Wood, Foxley Wood, and Wayland Wood contain only one of the species, Buff Wood has both. Woodell's (1969) data for ‘characteristic’ (i.e. single-species) populations at Marley and Lawn Woods are also included for comparison. n = number of plants in sample, S.E.M. = standard error of the mean. The ranges are inclusive (i.e. 20-60 = 20-00...—60-99...). due to a lower number of plants with very high (96-100%) stainability, especially in P. vulgaris. DISCUSSION MORPHOLOGICAL CHARACTERS AND INTROGRESSION P. elatior, P. Xpolyantha, and P. Xdigenea all have constant characters that allow them to be separated with certainty. Confusion between P. elatior and P. Xpolyantha probably arose amongst botanists of the 18th and 19th Century because few of them had ever seen P. elatior in the field, and they based their opinions upon dried specimens or written descriptions. Many of the useful characters (flower colour and markings, shape of the inflorescence, scent) are not preserved in herbarium specimens, and may not have been emphasised in the written descriptions. The main identification problem occurs in mixed populations of P. elatior, P. xdigenea, and P. vulgaris. Whilst there is no problem in identifying some hybrids (the classic “intermediate” types), or P. vulgaris, sepa- rating P. elatior from other hybrids becomes difficult for two reasons. First, P. elatior is rather variable in its hairiness, flower size and markings, and leaf shape and colour, more so it seems than native British P. vulgaris and P. veris, and some of the characters that are useful in distinguishing backcrossed or later generation hybrids from most Oxlips can be found in pure populations of P. elatior (e.g. cuneate leaves). Whether or not this variation, even in “pure” Oxlip populations, reflects historic introgression with P. vulgaris must remain an open question. However, it is of note that we did not find any populations where hybrids were detectable only through mol- ecular methods: the hybrids were all in woods with populations of Oxlips, Primroses, and obviously intermediate plants. Second, the amount of backcrossing and _ introgression means that some hybrids can be very close genetically and morphologically to P. elatior, and the boundaries of “Oxlip’, and “hybrid” become rather arbitrary and meaningless in hybrid swarms. Pollen fertility cannot be used to clarify the situation as it can in many other hybrid complexes, such as that involving Creeping Cinquefoil Potentilla reptans, Trailing Tormentil Potentilla anglica, and Tormentil Potentilla erecta (Harold 2006). The other potential pitfall is the rare Cowslip x Oxlip hybrid, P. xmedia. We have not been able to confirm this hybrid using molecular methods, and this would be a useful area of future work if plants could be found for analysis. We have found plants in mixed populations of P. veris and P. vulgaris that had obvious Cowslip characters, such as dark streaks on the petals, rather short hairs, and slightly inflated calyces, but these were tempered by longer hairs and larger and paler flowers than in P. veris. As either Oxlip or Primrose could have provided these features, determining which species is the other parent is more a matter of proximity than of morph- ology. 250 M. GURNEY, C. D. PRESTON, J. BARRETT & D. BRIGGS The presence of small flowers in a stalkless umbel in P. elatior is not associated with any sign of hybridisation (it is found in “pure” populations), and is part of the normal range of variation of the plant. It was noted by Miller Christy (1897), although he does not mention that the flowers were smaller than those on scapes. Miller Christy’s flowers were produced only early in the season, and we have seen them on plants before the normal inflorescences are produced. However, they are still in evidence later in the season, although they are much less obvious following the growth of the leaves and other vegetation, and it is possible that Miller Christy overlooked them. Tabor (1998) mentions a_ second flowering in some plants in June and July, in which the scape is very short. This differs from the sessile umbels that we have described in that the flowers are not carried on long pedicels (see photograph on page 97 of Tabor’s article), and we have not seen this ourselves, although similar looking inflorescences can rarely be found on plants during the normal flowering time. Other rare abnormalities we have found in P. elatior include the bracts being replaced by small leaves, the development of a second inflorescence lower down the scape, six- petalled flowers, flowers with two ovaries, and flowers with bright red stigmas (the last being a reasonably frequent phenomenon). EXTINCTION THROUGH HYBRIDISATION: MILLER CHRISTY’S HYPOTHESIS The view that in Buff Wood P. elatior seems to grade into P. xdigenea, whilst P. vulgaris remains a discreet entity could be seen as supporting Miller Christy’s (1897) suggestion that the “modest and retiring Oxlip is, in this country at least, being gradually hybridized out of existence by the more aggressive Primrose”’. However, this hypothesis has been rejected by Woodell (1969) and Rackham (1999). Our own investigations (Gurney 2000) found no reduction in fertility or seed production in Oxlips in Buff Wood when compared to those in “pure” populations, and we agree with Rackham that climate, woodland management, and herbivory are likely to be much more important in determining the species’ fate. ACKNOWLEDGMENTS Thanks to Hampshire County Council, the Wildlife Trusts, and the owners of the other sites for permission to work in_ their woodlands; to English Nature for consent to sample plants from Sites of Special Scientific Interest; to Martin Sanford for details of the Suffolk localities; Jeremie Fant for help in the fieldwork; Johannes Vogel, Steve Russell, Fred Rumsey, and Christina James of the Natural History Museum, and Steve Ansell and Nicky Williamson for help preparing the allozyme samples; Peter Walker for his data from Gamlingay Wood; and Roger Day and Dennis Farringdon for cultivating plants at the Department of Genetics Field Station, University of Cambridge. This work was supported by a NERC CASE studentship in partnership with CEH Monks. Wood. REFERENCES Brown, P. J. (1842). [Letter to Mr Edmonston]. The Phytologist 1: 192. BROWN, I. R., KENNEDY, D. & WILLIAMS, D. A (1982). The occurrence of natural hybrids between Betula pendula Roth and B. pubescens Ehrh. 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The Flora of Huntingdonshire and the Soke of Peterborough. Huntingdonshire Fauna and Flora Society, Peterborough. WOODELL, S. R. J. (1965). Natural hybridization between the Cowslip (Primula veris L.) and the Primrose (P. vulgaris Huds.) in Britain. Watsonia 6: 190-202. WOODELL, S. R. J. (1969). Natural hybridization in Britain between Primula vulgaris Huds. (the primrose) and P. elatior (L.) Hill (the oxlip). Watsonia 7: 115-127. (Accepted November 2006) —_ 4 to) i : “rire ie bo. } ey Le ' ‘ iS Watsonia 26: 253-269 (2007) LSS Which vascular plants are found at the northern or southern edges of their European range in the British Isles? C. D. PRESTON* CEH Monks Wood, Abbots Ripton, Huntingdon, Cambs. PE28 2LS ABSTRACT The northern and southern limits of the European distributions of native vascular plants in the British Isles are summarised. Of 1346 species considered, 279 reach their absolute northern limits in the British Isles and a further 167 at equivalent latitudes (45°- 60° N) elsewhere in Europe. A particularly high proportion of Mediterranean-Atlantic species in the British Isles reach their absolute northern limits here, and a higher proportion of Southern-temperate than Temperate species do so. By contrast, only 35 species are at their absolute southern limits in our area, with a further 7 extending south to equivalent latitudes (50°-60° N) elsewhere. The low southern limit totals reflect the presence of most northern species on high mountains in southern Europe. The range-edge species are listed, but the lists must be regarded as provisional as there are some difficulties in using the secondary sources on which they are based, the most serious being the problem of distinguishing native from introduced occurrences. Some 49% of threatened members of the British flora are at or near their northern or southern European limits in the British Isles. If Continental and Hyperoceanic species are also considered as species at the edge of their range, this figure increases to 64%. The increase is almost entirely due to the addition of many Continental species, which in Britain appear to be an especially threatened group. KEYWORDS: continental, distribution, major biome, Mediterranean-Atlantic, threat. INTRODUCTION Populations of plants at or near the edge of their distributional range are of great interest. In such situations plants may become increasingly restricted in their habitat require- ments, and these restrictions can _ offer considerable insights into the factors which limit the total distribution. Marginal populations (especially of annual species) may also fluctuate greatly in size from year to year, and this also provides evidence of ecologically limiting factors. They are sometimes genetically depauperate compared to core populations, or genetically adapted to specialised local environments. Marginal *e-mail: cdpr@ceh.ac.uk populations are especially interesting if the distribution of a species is changing, as they will often be the first ones to show the change. Global warming is widely anticipated in the next few decades and can, for example, be expected to result in a retreat of some species from the southern edge of their ranges accompanied by an expansion northwards. The conservation of species at the edge of their range has often been discussed, with some authors arguing that these populations should be safeguarded because of their intrinsic interest, whereas others caution against concentrating on the conservation of small, marginal populations of species which are more abundant elsewhere. In view of the general interest of edge-of- range populations, it is perhaps surprising that there is no easy way of identifying species which are at the edge of their range in the British Isles. This paper aims to fill this gap. Some species, especially those in_ the Mediterranean-Atlantic and Submediterranean- Subatlantic floristic elements such as Parietaria judaica, Polystichum setiferum (Fig. 1) and Umbilicus rupestris, have northern European limits which run diagonally from north-west to south-east Europe, and there is normally little doubt whether or not these reach their absolute northern limit in Britain. However, the northern limit of other species such as AHydrilla_ verticillata, Rumex conglomeratus and Salsola kali (Fig. 2) runs approximately west-east, and these may extend north to essentially the same latitude in several European countries. It is difficult to identify the precise location of their absolute northern limit, both because much of the available information is insufficiently precise geographically, and because the absolute limit may easily be extended by the discovery of new populations. In this paper, therefore, I have listed the species which reach their northern and _ southern European limits as native species either in the British Isles itself or at similar latitudes else- where in Europe. 254 C. D. PRESTON 30 2702 eno) fo ae St f O a 7. ae soot at i ~ Me oy 12° 18° 24° a 36° 42° 48° 54°60° 576621 TS \ Niet F \ ~ \ \ eee te eet ee FIGURE 1. The distribution of Polystichum setiferum in Europe (Jalas & Suominen 1972). This Submediterranean-Subatlantic species has a northern limit running NW-SE and is clearly at its northern European limit in the British Isles. IDENTIFYING THE EUROPEAN RANGES OF BRITISH AND IRISH SPECIES SPECIES CONSIDERED My aim has been to assess the European distributional limits of all the native species of the British Isles, with the exception of the microspecies of Hieracium, Rubus and Tarax- acum. Species which are endemic to the British Isles or are confined in Europe to our area are excluded for the obvious reason that they have no wider European. distribution, and infraspecific taxa have not been considered. Taxonomy and nomenclature usually follow Stace (1997) and the species considered are those treated by Preston, Pearman & Dines (2002) as native or doubtfully native (‘native or alien’). In a few cases I have departed from these sources to take account of recently published work or to exclude species with distributions which are not sufficiently well understood for their limits to be assessed. Asparagus officinalis subsp. prostratus 1s treated as a species, A. prostratus, following Kay, Davies & Rich (2001), and two species added to the flora since 1997, the native Carex salina (Dean et al. 2005) and the apparently native Cystopteris diaphana (Murphy & Rumsey 2005), have been _ considered. Cystopteris alpina has been restored to the British list by Rumsey (2003), although it is now believed to be extinct. Segregates in the Callitriche hamulata, Arctium minus, Erophila verna, Salicornia europaea, S. procumbens and Utricularia intermedia complexes and_ the species Carex recta, Cystopteris dickieana, Dactylorhiza lapponica, D. majalis and D. traunsteineri have not been considered because PLANTS WITH RANGE EDGES IN THE BRITISH ISLES 255 40° ¢ [BS e i, Tee ° 12° 18° 24° 30° 36° 42° 48°~54°S 60°=2506") 72°C 65° | paltenl | ae aN 7° nN 4 SD yay Pre pl K —E Ee ) Na SF f y é X 7 Ne ro) ° =~, | ra \ t \ \ > \ iN ae \ fe > \ > \ ht i K, , \ ) sk 5 FIGURE 2. The distribution of Salsola kali in Europe (Jalas & Suominen 1980). The symbols ‘x’ and ‘+’ denote ‘probably extinct’ and ‘extinct’ respectively, and occurrences as an established alien are shown by open circles. This Eurosiberian Southern-temperate species reaches its northern limit at similar latitudes in Scotland, Norway, Sweden, Finland and Russia. of doubts about their taxonomic status, distributional uncertainties or because recent taxonomic revisions have reduced them to endemic status. Viola odorata and _ the doubtfully native Onobrychis viciifolia and Ribes rubrum have also been excluded as their native ranges are so confused by introductions that the northern and southern limits cannot be assigned to the latitudinal bands specified below. Valerianella eriocarpa, considered a neophyte by Preston et al. (2002), is treated as doubtfully native following a reassessment of its status (Pearman & Edwards 2002). Symphytum tuberosum is also treated as doubt- fully native; it was considered native, with reservations, by Preston et al. (2002) but, as Braithwaite, Ellis & Preston (2006) suggest, it is “much more likely to be a neophyte”. SOURCES OF DISTRIBUTIONAL DATA In compiling information on the distributional limits of species, I have drawn on published distribution maps, especially those of Atlas Florae Europaeae (Jalas & Suominen 1972- 1994; Jalas, Suominen & Lampinen 1996; Jalas et al. 1999; Kurtto, Lampinen & Junikka 2004) or, for the majority of taxa not yet covered by this Atlas, the maps compiled by Hultén & Fries (1986). For plants covered in neither of these works, I have consulted Meusel, Jager & Weinert (1965), Meusel et al. (1978) and Meusel & Jager (1992) for central European species, and Bolos & Vigo (1984-2001), who map many southern European species, as well as Tutin et al. (1968-1980, 1993). For more detailed information on the distribution of species in countries in the same latitudinal 256 C. D. PRESTON TABLE 1. SUMMARY OF MAJOR BIOME CATEGORIES (PRESTON & HILL 1997) Name Arctic-montane Boreo-arctic Montane Wide-boreal Boreal-montane Boreo-temperate Wide-temperate Temperate Southern-temperate Mediterranean-Atlantic Explanation Main distribution in the tundra or above the tree-line in temperate mountains. In both the tundra and the coniferous forest zones. Extends from the temperate zone to the tundra. Main distribution in the coniferous forest zone. In both the coniferous and the broadleaf forest zones. Extends from the Mediterranean zone to the coniferous forest zone. Main distribution in the broadleaf forest zone. In both the Mediterranean region and the broadleaf forest zones. In the Mediterranean region, and extending north into the broadleaf forest zone in western Europe. Submediterranean-Subatlantic and Mediterranean-montane species are included here. band as the British Isles, useful sources include Hultén (1971), Mossberg & Stenberg (2003) and Jonsell (2000, 2001) for the Nordic countries, Mennema, Quené-Boterenbrood & Plate (1980, 1985) and van der Meijden, Plate & Weeda (1989) for the Netherlands, Haeupler & Schonfelder (1988) and Benkert, Fukarek & Korsch (1996) for Germany, Institut Floristique Franco-Belge (1978-2001) for Belgium and northern France and Dupont (1962) for species of the Atlantic zone. The native ranges of species in Britain are taken from Preston et al. (2002), with additional information from local floras such as Graham (1988), Scott & Palmer (1987) and Swan (1993). Distributional limits are analysed below in relation to the major biome categories defined by Preston & Hill (1997), Preston et al. (2002) and Hill, Preston & Roy (2004) (Table 1). Of the three species recently added to the flora, Carex salina and Cystopteris alpina have been treated as European Boreo-arctic Montane and C. diaphana as Suboceanic Southern- temperate. CLASSIFICATION OF NORTHERN AND SOUTHERN LIMITS The above sources were used to classify the northern European limit of the species considered into five latitudinal bands: 45°-50° N, 50°-55° N, 55°-60° N, 60°-65° N and >65° N. Similarly, the southern limits of species were classified into the bands <45° N, 45°-50° N, 50°-55° N and 55°-60° N. Almost all the land area of the British Isles falls within the 50°-55° and 55°-60° bands (Figs 1-2). The only parts of the British Isles south of 50° N are the Channel Islands, the Isles of Scilly and the southern part of the Lizard peninsula; Shetland alone extends north of 60° N. (None of the species considered has a southern limit in the 60°—65° band, which would imply growing as a native in Shetland but descending no further south in Britain.) The 55° N line of latitude very approximately separates Ireland, Wales and England to the south from Scotland to the north. Thus plants occurring as natives in the British Isles and reaching their northern limits in Europe between 45° and 60° are usually close to their northern limits here, even if they do not reach their absolute European limit in these islands. Similarly most plants extending south to latitudes between 50° and 60° are close to their southern limits in Britain. Species extending south to 45°-50° are not necessarily close to their southern limits in the British Isles as they may extend almost 5° further south; by contrast, a native of the British Isles which reaches its northern limit between 45° and 50° must be near its limit in our area. In addition to allocating the limits of species to these latitudinal bands, I have tried to identify the plants reaching their absolute northern and southern limits in our area. In determining species limits, all recorded native sites are considered, including those at which the species has become extinct in recent times. The data on _ distributional limits are summarised in this paper, and species at or near their northern or southern limits in the British Isles are listed in Appendix | and 2. The full data have been added to the PLANTATT database (Hill et al. 2004) and can be downloaded from the Biological Records Centre website (www.brc.ac.uk). PLANTS WITH RANGE EDGES IN THE BRITISH ISLES Zie)i} TABLE 2. NORTHERN EUROPEAN LIMITS OF NATIVE SPECIES IN RELATION TO THEIR OCCURRENCE IN MAJOR BIOMES Major biome Number of species at N limit in latitudinal band Total 45°—50° 50°—55° 55°—60° 60°—65° >65° Arctic-montane JH Bye TS 4*+75 Boreo-arctic Montane je 1 39 1*+40 Wide-boreal 19 19 Boreal-montane qe 5 86 7*+91 Boreo-temperate jeep 8 DAD 1*+221 Wide-temperate 38 3)3) Temperate 2*+1 30*+11 39*+82 5*+192 141 76*+427 Southern-temperate Sel 43*+19 37*+38 a0) Sy 88*+145 Mediterranean-Atlantic 3*+] 67*+2 31*+11 ] 101*+15 Unknown I% 1 1*+1 Total 8*+3 141*+32 120*+132 10*+257 643 279*+1067 *The askerisked figures are the number of British species at their absolute northern limit in the British Isles. RANGE LIMITS IN RELATION TO OCCURRENCE IN MAJOR BIOMES The northern limits of native species are summarised in Table 2. In all 279 of the 1346 species considered (21%) reach their absolute northern limits in the British Isles, and a further 167 (12%) extend north to equivalent latitudes (45°—60° N) in mainland Europe. These species are listed in Appendix 1. Almost all the species of northern biomes, and the widespread species, extend to latitudes north of 65°. A small number of Arctic-montane and Boreal- montane species occur at the northern edge of their range in Britain, these being Alpine species which are absent from the Arctic, e.g. the Arctic-montane Minuartia recurva, M. sedoides and Myosotis alpestris, the doubtfully native Boreo-arctic Montane Homogyne alpina and the Boreal-montane Meum athamanticum, Oxytropis halleri and Viola lutea. Thlaspi caerulescens extends north to Scotland as a native, but is very widely naturalised in the Nordic area (Jalas et al. 1996) and Myosotis stolonifera occurs in the mountains of the Iberian peninsula and the hills of northern England and southern Scotland. Plants of the Temperate and Southern biomes show an obvious tendency to reach their northern limits further south. Of the species reaching their northern European limits between 45° and 60° N, there is a clear increase in the percentage occurring at their absolute limit in Britain from Temperate (43%) through Southern-temperate (59%) to the Mediterranean-Atlantic (88%) major biome categories. This presumably reflects the fact that Temperate species may be able to extend further north in mainland Europe because of the warmer continental summers, whereas the Mediterranean-Atlantic species require milder winters and are therefore likely to extend further north in the oceanic west of Europe. The difference between Temperate and Southern-temperate species on the one hand and Mediterranean-Atlantic species on the other is almost inevitable in view of the definition of the major biome categories, as the Mediterranean-Atlantic plants differ from the Temperate and Southern-temperate species in occurring further north in the Atlantic zone of western Europe than they do further east. There is a rather unexpected trend for a greater proportion of species reaching their northern limit between 50° and 55° N in Europe to have their absolute limit in the British Isles compared to those which extend north into the 55°—60° N band. In the southerly latitudinal band 82% of species reach their northern limits in the British Isles, compared to 48% of those in the northerly band, a highly significant difference (p<0-001). This difference holds within all three of the predominant major biome categories. For Temperate species 73% of the 50°—55° species are at their northern limit here compared with 32% for the 55°-60° plants (p<0-001); the equivalent figures for Southern-temperate species are 69% vs 49% (p<0:05) and for Mediterranean-Atlantic species 97% vs 74% (p<0-001). Most of the (rather few) species in 258 C. D. PRESTON the 50°-55° zone which reach their absolute limits outside the British Isles extend further north in Germany. The larger group reaching their northern limits outside our area in the 55°-60° zone include many at their northern limits in Denmark or Sweden, smaller numbers in Norway or in the eastern Baltic (including the Baltic states and Baltic Russia) and very few species reaching their limits further east in Russia. The reasons for this shift in northern limits to continental Europe for the more northerly species are not entirely obvious, and the fact that many species have become naturalised after they have been introduced to sites north of their native limits suggests that these reasons are unlikely to be simple. Factors to be considered include differences in the proportion of the total land area in each latitudinal band which lies in the British Isles and the range of climates and habitats provided in southern Britain and Ireland compared to northern Britain. The contrast between the climate of S.W. Ireland and East Anglia in the 50°-55° zone is much greater than the climatic extremes within the northern zone. Another factor is almost certainly the paucity of lowland, calcareous habitats in much of Scotland. Ellenberg R values, which indicate pH preferences from extreme calcifuge (R = 1) to calcicole (R = 9, see Hill et al. 2004) can been used to identify the number of marked calcicoles at their absolute limits in Britain and elsewhere in Europe (Table 3). A few species with R = 8-9 are halophytes rather than calcicoles but they do not greatly affect the percentages in Table 3. The overall proportion of calcicoles is higher for the species with northern limits between 50° and 55° than for those with limits between 55° and 60° N (p<0:05), not unexpectedly in view of the fact that acidic habitats predominate in the Boreal zone. More relevantly, the proportion of calcicoles amongst the species reaching their absolute limits in Britain is significantly lower in the northerly band than in the southerly band (p<0-001). The calcicoles occurring at their northern limits between 55° and 60° N in continental Europe include a small but well- known group of species which extend north to the calcareous islands of Oland and Gotland in the Baltic (e.g. Aster linosyris, Helianthemum oelandicum, Orchis laxiflora, Potamogeton coloratus). Whereas there is a clear relationship between major biome categories and northern limits, there is no such relationship with southern limits (Table 4). The majority of species in all major biome categories extend south to southern Europe (<45° N). This reflects the presence of high mountains at southern latitudes in Europe, and hence the availability of habitats above the tree-line. A few Arctic and Boreal species are absent from the south- ern European mountains, and these represent the majority of species which attain their southern limits in the British Isles. Only 35 species are at their absolute southern limits in our area and a further 7 extend south to equivalent latitudes (50°-60°) in mainland Europe (Appendix 2). The high proportion of these species which are at their absolute southern limit in the British Isles is noteworthy, and indeed all 15 species extending south to between 55° and 60° are at their absolute limit here. There is in fact little high ground in Europe between 50° and 60°, so montane species which extend south to the British Isles but not to the mountains of central Europe are likely to be at their absolute southern limits here. ARE THREATENED PLANTS IN BRITAIN MAINLY PLANTS AT THE EDGE OF THEIR RANGE? IUCN threat categories for British species have recently been revised (Cheffings & Farrell 2005). Table 5 summarises the number of threatened species in Britain which are close to their northern or southern limits. Only the species covered in this paper are included in Table 5; other Red Listed species (notably archaeophytes, endemics, plants confined to the British Isles in Europe and _ infraspecific taxa) are excluded from consideration. I have used the term ‘threatened species’ to include those classified as Extinct, Extinct in the Wild, Critically Endangered, Endangered, Vulnerable and Near Threatened in the Red List. Of the threatened species considered, almost half (49%) are at or near their northern or southern limits. Although this is a higher percentage than the equivalent figure (36%) for species in our flora as a whole, it is clear that many species at the limits of their latitudinal range are not threatened, and that many threatened species have a very broad latitudinal range in Europe. It might be argued that some threatened (and non-threatened) species are at their western and eastern limits in our area. In general, western limits are of little interest to British and Irish PLANTS WITH RANGE EDGES IN THE BRITISH ISLES 2359 TABLE 3. ELLENBERG R VALUES OF SPECIES REACHING THEIR NORTHERN EUROPEAN LIMITS BETWEEN 50° AND 60° N Northern limit No. of species Ellenberg R values 50°—55° N, absolute limit in Britain 1407 50°-55° N, absolute limit elsewhere 3) 55°-60° N, absolute limit in Britain 120 55°-60° N, absolute limit elsewhere 1317 Range Mean % calcicoles (R = 8-9) 1-9 6:4 31 4-9 6:8 44 2-9 6:0 12 2-9 6:7 30 + These totals each exclude one species for which an Ellenberg R value has not been allocated. TABLE 4. SOUTHERN EUROPEAN LIMITS OF NATIVE SPECIES IN RELATION TO THEIR OCCURRENCE IN MAJOR BIOMES Major biome Number of species at S limit in latitudinal band Total <45° 45°-S0° 50°-55° 55°-60° Arctic-montane 33 7 5*+1 Bi 18*+61 Boreo-arctic Montane 29 3 6*+2 i 7*+34 Wide-boreal 17 yy 19 Boreal-montane 78 8 8*+3 il* 9*+89 Boreo-temperate 216 1*+5 1*+221 Wide-temperate 38 333) Temperate 49] 12 503 Southern-temperate Day 1 233 Mediterranean- Atlantic 116 116 Unknown 1 1 3 Total 1266 1*+38 19*+7 > 35*+1311 *The asterisked figures are the number of British species at their absolute southern limit in the British Isles. botanists, as the British Isles are at the western fringe of Europe. All our species approach their western limits here. Whether or not one of our common species reaches its absolute western European limit in the British Isles therefore depends on whether it occurs on the Atlantic islands of Iceland and the Azores (the only parts of Europe as defined by Flora Europaea that are further west than western Ireland) and on the exact details of its distribution in the British Isles, western France and the Iberian peninsula. Although the details of its distri- bution in the western fringes will be determined by its ecology, the relevant eco- logical factors will include _ dispersal mechanisms and habitat preferences which are unlikely to act together to make “presence at its absolute western limit in Britain’ a useful piece of information about the species. Species which reach a western European limit in eastern Britain are perhaps more clearly at their western limit here, and the more extreme examples are identified as ‘Continental’ species by Preston & Hill (1997). One small group of species, the ‘Hyperoceanic’ species, have such western distributions in Europe that they are close to their eastern limits in Britain, and these are listed by Preston & Hill (1999). If the Continental and Hyperoceanic species as well as those close to their northern and southern limits are considered, the percentage of threatened species which are at or near the edge of their range increases from 49% to 64%. This is almost entirely due to the addition of | substantial number of Continental species which have a broad latitudinal range but are threatened in Britain. Only a single threatened Hyperoceanic plant is not at either its northern C. D. PRESTON 260 aU) UL (GOT) HELP -suosval JoyjO Joy Joded sty) ul UONeJopIs sSuryyouD Aq popnpour osoMm “UIeILIE JID BptsIno Sofs] YSU © “s]B]0} JILL] UOYINOS ot) JOU Ng UOYWOU oy} UTP UJOYINOS PUL UJOYOU Sj YO SoYoVaA DID]]}ONAIA DIPAPAH “S~eIOWNS IIe Sore apnjoul st U0d Woy popnyoxe aavy J Soloeds oy) IssuOUTR ore JO AST] SUIIL AA, y} Jo sued osou) 0} PoyoENsod o1e Koy) Sv Patfisse[o Jou o19M sotsods osoY Lx. ‘Zununod-s[qnop ploar OL ‘odoing puvyureu Ul JI UL SULUNJOS puR SMOY “AJOBOILO IST] POY YRS UL STRIO) dy) 0} aJvJor SoseyUsoIJOd OY], 209 PUR ,QS UdEMIJOq SITU] 9c l (%6S) 06L (%Ib) 9SS (%S) 69 (%1>) 9 (%€) SE (%T1) LOI (%17) 6L7 [RIOL (Ge (%91) ¢ (%b8) LZ (%7T) L (%79) OC 4: POTJISSB[D JON Ul (%LT) € (%EL) 8 (%9b) S (%LT) € JUSTOTJOP BI 6001 (%L9) 9L9 (%EE) EEE (%€) 9C (%I>) € (%7) LI (%01) OOI (%81) L8I UIQUOD ]SP9"] VOC (%9€) 901 (%F9) 881 (%S1) EF (%1) € (6P)ET (%17) 09 (%r7) 69 pauajpad yf [DIOL fe, (%CP) CE (%8S) Sr (%01) 8 (%1) 1 (%v) € (%81) vi (%S7) 61 pousyvolY} IGON Del (%8E) 1S (%79) C8 (%v1) 61 (%9) 8 (%91) TT (%9T) OE a[qeiou[nA cS (%7¢) LI (%89) GE (%61) OI (%T) I (%b) T (%97) V1 (%L1) 6 pososuepud 91 (%9) | (%F6) SI (%1€) S (%bvr) L (%61) € pososuepus ATRON) Ell (%Tv) S (8S) L (%8) | (%8) | (%S7) € Cyne. younx| [eyuouTjUuOS 10U soroods odoin odoing ul oruvasosodAy Joyyiou. §—sy#uuyy aSuUDA — FeYUOUTJUOD JOU MIYMES|I N 09 d19YMOS]I N .09 pue odoing Ut ,Q¢ Jo ADAU AO otuvas010dAy pur ,0¢ uaomiog = SOS YSU UL pure CP usoMjoq = SIS] YSHIIG Ur yynos pue ,99 Jo you = JD saioads jeuonippe JWT] UJOYINOS JT] UIOYINOS } UT] UJOY}IOU J UT] UOUIIOU [R10], Burpus}xo sotoads “ON fo ‘ou [DIOL jo ‘ON ye sorseds ‘ON ye soroods ‘ON ye sotoods “ON ye soroods ‘ON AJOBIIe9 IST] Poa (CQOT TIHMAVA 2 SONISHAHD) KYOOALVO LSIT GAY NONI MAHL OL NOLLV Tada NI SqIodds AALLVN AO SLIAIT NVadOund *s aTaVL PLANTS WITH RANGE EDGES IN THE BRITISH ISLES 261 or southern limit, the Near Threatened Hymenophyllum wilsonii. This highlights the threat to Continental species in Britain, a group first discussed in detail by Walters (1953). Of the 88 species which are listed by Preston & Hill (1997) as Continental and considered in this paper, 86 occur in Great Britain and of these 54 (63%) are listed as Extinct (2), Critically Endangered (7), Endangered (12), Vulnerable (22) or Near Threatened (11) by Cheffings & Farrell (2005). DISCUSSION LIMITATIONS OF THE SOURCES The user of secondary sources to identify the limits of the native range of species in Europe is confronted by three major problems. Was the information presented taxonomically accurate at the time of publication, and has it been superseded by subsequent taxonomic revision? Are native records distinguished from alien (including casual) occurrences? Do sources such as small-scale distribution maps provide sufficient geographical precision? All these are serious problems. To a great extent, however, users are in the hands of their sources. It is clearly impractical to investigate the European distribution of all native British species individually, and even if one did there is no likelihood that the result would be any better than the published compilations. By maintaining a semi-critical approach to the sources one can attempt to identify potential problems and check one source against another. Outlying localities on small-scale European maps can be checked against national floras and atlases, for example. These publications can also be used to provide more precise details of the limits of species than are apparent on small-scale maps. More critical accounts of the distribution of some species are available in monographic works. The greatest problem is the identification of native ranges. Even if the native range is known to the local observer, it may not be clear to a compiler working elsewhere in Europe or the compiler may choose not to distinguish alien occur- ences on the map. Outlying northern localities on many of the maps compiled by Hultén & Fries (1986), for example, represent introduced or even casual occurrences. In the cases of Scandinavian records, these maps can be checked against Mossberg & Stenberg’s (2003) maps and national and _ regional floras. However, there are limits to the number of such checks which are either possible or practical in a reasonable time-scale. The interpretation of the native ranges of British and Irish species is that of Preston et al. (2002). The problems associated with assessing native ranges in Britain, which to a large extent reflect those associated with the wider problem in Europe as a whole, are discussed briefly on pp. 10-11 of that work. For the above reasons, the lists provided in this paper are provisional. I hope _ that specialists in individual species or genera will be able to review the assessments and correct them where necessary. The lists will also need revision in the light of the critical reviews of the distribution and native or introduced status of Nordic plants which will become available as further volumes of Flora Nordica are published. POTENTIAL USEFULNESS OF INFORMATION ON DISTRIBUTIONAL LIMITS This paper has been prepared with several potential uses in mind. Authors of local Floras can use the lists in Appendix | and 2 in conjunction with the maps in Preston ef al. (2002) to identify those species at their northern or southern limits in their area. The lists could also be used in planning scientific studies that compare populations of species at the edge of their range with those in a more central position. The information on individual species may also be useful in assessing the extent to which edge-of-range populations are conserved in protected sites. However, one of the main aims of this compilation is to provide a further ‘plant attribute’ to be used in interpreting the observed changes in the British and Irish flora. Even a simplified version of Preston & Hill’s (1997) major biome categories correlated surprisingly well with the changes detected in several Broad Habitats by the BSBI ‘Local change’ survey (Braithwaite et al. 2006). Information on the northern limits is potentially useful, although it may not always prove to be as informative as the classification by major biome categories. The latter summarise the overall range of a species and disregard outlying occurrences, whereas by definition the distributional limits are governed by the outlying occurrences. At outlying localities a population may be dependant on a _ very specialised microhabitat, or the plants may be relictual, dating from a different climatic 262 C. D. PRESTON period. Although such relict populations persist they may, in the extreme case of some long- lived species, no longer even be capable of sexual reproduction in the current climatic regime. Once maps such as those published in Atlas Florae Europaeae are available for all species it will be possible to use numerical methods to summarise range boundaries, with rules to determine whether or not disjunct localities should be treated as outliers, but at present such maps are available for only a minority of European species. Information on the southern limits of species is much less informative than that on northern limits as, unlike major biome categories, it fails to take into account the altitudinal range of species. The current interest in the effects of climate change has led to an increasing concentration of research on the behaviour of populations at the edge of their range. It 1s clear from the results presented here that British and Irish botanists are ideally placed to study the behaviour in the years to come of species at the northern edge of their European ranges. This is surely an area in which the B.S.B.I. could develop one of those “really impressive examples of botanical co-operation in which the amateur and professional talent of our Society is used with great effect” (Walters OWS): ACKNOWLEDGMENTS I began to investigate this topic when working with David Pearman on a contract to report on the European ranges of British species to the Countryside Council for Wales. I am grateful to Andy Jones of CCW for providing this opportunity, to David Pearman for extracting information from the publications of Institut Floristique Franco-Belge and to Clive Jermy (Carex), Thomas Karlsson (Nordic plants), Hildemar Scholz (Agrostis) and Clive Stace (Agrostis, Festuca) for providing detailed information on the distribution of particular species. I thank Peter Rothery for useful discussion and Mark Hill, David Pearman and Kevin Walker for valuable comments on a draft of the paper. I thought it appropriate to complete the paper for the issue of Watsonia dedicated to Dr S. M. Walters as it examines the British flora in a European perspective. Like many others, I got much encouragement from Max in my early years as a botanist, and in particular I was influenced by a brief address he gave to the Cambridge Natural History Society when I was an undergraduate. This so captured the romance of field botany that it left me in no doubt that I should pursue an interest in plants rather than a hitherto much stronger interest in entomology. REFERENCES BENKERT, D., FUKAREK, F. & KORSCH, H., eds (1996). Verbreitungsatlas der Farn- und Bliitenpflanzen Ostdeutschlands. Jena: Gustav Fischer. BOLOS, O. DE & VIGO, J. (1984-2001). Flora dels Paisos Catalans. Volume 1 (1984), 2 (1990), 3 (1995), 4 (2001). Barcelona: Editorial Barcino. 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Huntingdon: Centre for Ecology and Hydrology. HULTEN, E. (1971). Atlas 6ver vdxternas utbredning i Norden. Stockholm: Generalstabens Litografiska Anstalts Forlag. HULTEN, E. & FRIES, M. (1986). Atlas of north European vascular plants north of the Tropic of Cancer. 3 vols. K6nigstein: Koeltz Scientific Books. INSTITUT FLORISTIQUE FRANCO-BELGE (1978-2001). Documents floristiques 1-6. Bailleul: Centre Regional de Phytosociologie. , JALAS, J. & SUOMINEN, J., eds (1972-1994). Atlas Florae Europaeae. Volume 1 (1972), 2 (1973), 3 (1976), 4 (1979), 5 (1980), 6 (1983), 7 (1986), 8 (1989), 9 (1991), 10 (1994). Helsinki: Committee for Mapping the Flora of Europe and Societas Biologica Fennica Vanamo. PLANTS WITH RANGE EDGES IN THE BRITISH ISLES 263 JALAS, J., SUOMINEN, J. & LAMPINEN, R., eds (1996). Atlas Florae Europaeae. Volume 11. Helsinki: Committee for Mapping the Flora of Europe and Societas Biologica Fennica Vanamo. JALAS, J., SUOMINEN, J., LAMPINEN, R. & KURTTO, A., eds (1999). 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The Hague: Dr. W. Junk. MENNEMA, J., QUENE-BOTERENBROOD, A. J. & PLATE, C. L., eds (1985). Atlas van de Netherlandse Flora. 2 Zeldzame en vrij zeldzame planten. Utrecht: Bohn, Scheltema & Holkema. MEUSEL, H. & JAGER, E. J. (1992). Vergleichende Chorologie der zentraleuropdischen Flora. Volume 3. 2 vols. Jena: Gustav Fischer. MEUSEL, H., JAGER, E., RAUSCHERT, S. & WEINERT, E. (1978). Vergleichende Chorologie der zentraleuropdischen Flora. Volume 2.2 vols. Jena: Gustav Fischer. MEUSEL, H., JAGER, E. & WEINERT, E. (1965). Vergleichende Chorologie der zentraleuropdischen Flora. Volume |. 2 vols. Jena: Gustav Fischer. MOSSBERG, B. & STENBERG, L. (2003). Die nya nordiska Floran. Wahlstro6m & Widstrand. MURPHY, R. J. & RUMSEY, F. (2005). Cystopteris diaphana (Bory) Blasdell (Woodsiaceae) — an overlooked native new to the British Isles? Watsonia 25: 255-263. PEARMAN, D. A. & EDWARDS, B. (2002). Valerianella eriocarpa Desv. in Dorset, and a reassessment of its status as a presumed introduction in Britain. Watsonia 24: 81-89. PRESTON, C. D. & HILL, M. O. (1997). The geographical relationships of British and Irish vascular plants. Botanical Journal of the Linnean Society 124: 1-120. PRESTON, C. D. & HILL, M. O. (1999). The geographical relationships of the British and Irish flora: a comparison of pteridophytes, flowering plants, liverworts and mosses. Journal of Biogeography 26: 629- 642. PRESTON, C. D., PEARMAN, D. A. & DINES, T. D. (2002). New atlas of the British and Irish flora. Oxford: Oxford University Press. RUMSEY, F. (2003). Bladder-ferns Cystopteris of the British Isles. British Wildlife 15: 93-96. SCOTT, W. & PALMER, R. (1987). The flowering plants and ferns of the Shetland Islands. Lerwick: Shetland Times. STACE, C. A. (1997). New Flora of the British Isles, edn 2. Cambridge: Cambridge University Press. SWAN, G. A. (1993). Flora of Northumberland. Newcastle upon Tyne: Natural History Society of Northumbria. TUTIN, T. G., BURGES, N. A., CHATER, A. O., EDMONDSON, J. R., HEYWOOD, V. H., MooreE, D. M., VALENTINE, D. H., WALTERS, S. M. & WEBB, D. A., eds (1993). Flora Europaea. Volume 1, edn 2. Cambridge: Cambridge University Press. TUTIN, T. G., HEYWOOD, V. H., BURGES, N. A., MOORE, D. M., VALENTINE, D. H., WALTERS, S. M. & WEBB, D. A., eds (1968-1980). Flora Buropaea: Volume 2 (1968), 3 (1972), 4 (1976), 5 (1980). Cambridge: Cambridge University Press. WALTERS, S. M. (1953). The continental element in our flora, in The changing flora of Britain (ed. J. E. Lousley), pp. 124-129. Botanical Society of the British Isles, Oxford. WALTERS, S. M. (1975). Presidential address 1974: Botanical co-operation. Watsonia 10: 325-330. (Accepted September 2006) 264 C. D. PRESTON APPENDIX |. SPECIES AT OR NEAR THE NORTHERN EDGE OF THEIR NATIVE EUROPEAN RANGE IN THE BRITISH ISLES Native British species which have their northern European limits in the latitudinal bands 45—50° N, 50-55° N and 55-—60° N are listed in this section. Species are asterisked if the absolute limit is reached in the British Isles, but not if it they reach their northern limit at the same latitude in our area and in other European countries or if the precise northern limit is obscured by introductions. The following additional annotations are used: § Doubtfully native (1) Confined as a native in the British Isles to the Channel Islands (2) Confined as a native in the British Isles to Ireland (3) Limit(s) of native range particularly confused by introductions (this also applies to doubtfully native taxa) SPECIES WITH NORTHERN EUROPEAN LIMITS BETWEEN 45° N AND 50° N Anogramma leptophylla* (1) Armeria arenaria (1) Exaculum pusillum* (1) Festuca armoricana (1) Festuca huonii* (1) Limonium auriculae-ursifolium* (1) Limonium normannicum* (1) Myosotis sicula* (1) Ophioglossum lusitanicum* Ornithopus pinnatus* Ranunculus paludosus (1) SPECIES WITH NORTHERN EUROPEAN LIMITS BETWEEN 50° N AND 55° N Aceras anthropophorum* Aconitum napellus§* Adiantum capillus-veneris* Agrostis curtisii* Ajuga chamaepitys§ Allium sphaerocephalon§ Alopecurus bulbosus Anagallis arvensis (3) Arabis scabra* Arbutus unedo* (2) Arum italicum* Asparagus prostratus Asperula cynanchica* Asplenium onopteris* (2) Atropa belladonna* (3) Brassica oleracea§* Bromopsis erecta* Bunium bulbocastanum Bupleurum baldense* Buxus sempervirens* (3) Callitriche truncata* Carex humilis Centaurium scilloides* Centaurium tenuiflorum* Cirsium eriophorum* Cirsium tuberosum Clematis vitalba (3) Clinopodium ascendens* Clinopodium calamintha* Clinopodium menthifolium Colchicum autumnale* Corrigiola litoralis Crassula tillaea* Cynodon dactylon§ Cynoglossum germanicum* Cyperus longus* Cystopteris diaphana* Daboecia cantabrica* (2) Damasonium alisma* Daphne laureola* Dianthus gratianopolitanus Draba aizoides* Epilobium lanceolatum* (3) Erica ciliaris* Erica erigena* (2) Erica vagans* Erodium maritimum* Euphorbia amygdaloides* Euphorbia peplis* Euphorbia serrulata§ Festuca longifolia* Frankenia laevis* Gagea bohemica Galium constrictum* Galium parisiense§* Gastridium ventricosum§* Gaudinia fragilis§* Gentianella germanica Geranium purpureum* Geranium rotundifolium* Gladiolus illyricus* Gnaphalium luteoalbum§ Helianthemum apenninum* Helleborus foetidus* Helleborus viridis§* Herniaria ciliolata* Himantoglossum hircinum* Hippocrepis comosa* PLANTS WITH RANGE EDGES IN THE BRITISH ISLES Hypericum linariifolium* Hypericum undulatum* Iberis amara* Inula crithmoides* Iris foetidissima* Isoetes histrix* Juncus acutus* Koeleria vallesiana* Lactuca saligna* Lathyrus aphaca§* Lathyrus nissolia* Leucojum aestivum* Limonium bellidifolium* Limonium binervosum* Linum bienne* Lithospermum purpureocaeruleum* Lobelia urens* Lotus angustissimus Lotus subbiflorus* Ludwigia palustris Luzula forsteri* Matthiola sinuata§* Medicago arabica* Medicago polymorpha* Melittis melissophyllum Mentha pulegium* Mentha suaveolens* (3) Mibora minima* Milium vernale (1) Minuartia hybrida* Minuartia recurva* (2) Muscari neglectum§* Neotinea maculata* Oenanthe pimpinelloides* Oenanthe silaifolia* Ononis reclinata* Ophrys fuciflora Ophrys sphegodes* Orchis simia* Ornithogalum pyrenaicum* Otanthus maritimus* Petrorhagia nanteuilii* Petroselinum segetum* Peucedanum officinale Physospermum cornubiense Pinguicula grandiflora* (2) Poa infirma* Polycarpon tetraphyllum§* Polygala calcarea* Polygonum maritimum* Polypogon monspeliensis* Potamogeton nodosus Puccinellia fasciculata* Pulmonaria longifolia* Pyrus cordata§* Ranunculus parviflorus* Ranunculus tripartitus Romulea columnae* Rosa stylosa* Rubia peregrina* Rumex pulcher* Rumex rupestris* Ruscus aculeatus* Salicornia pusilla* Saxifraga hirsuta* (2) Schoenoplectus pungens§ Schoenoplectus triqueter Scilla autumnalis* Scirpoides holoschoenus Sedum forsterianum* Serapias parviflora §* Sibthorpia europaea* Simethis planifolia* (2) Sison amomum* Sorbus aria* Sorbus domestica* Spartina maritima* Spiranthes aestivalis Stachys germanica* Suaeda vera* Tamus communis* Thesium humifusum* Thlaspi perfoliatum Trifolium bocconei* Trifolium glomeratum* Trifolium incarnatum * Trifolium occidentale* Trifolium ochroleucon* Trifolium ornithopodioides* Trifolium squamosum* Trifolium strictum* Trifolium subterraneum* Trifolium suffocatum* Trinia glauca* Tuberaria guttata* Ulex minor* Ulmus proceras* Valerianella eriocarpa§s* Verbascum pulverulentum§* Viburnum lantana* Vicia bithynica* Vicia parviflora* Viola kitaibeliana Viola lactea* Vulpia ciliata* 265 266 SPECIES WITH NORTHERN EUROPEAN LIMITS BETWEEN 55° N AND 60° N Acer campestre* Alisma gramineum Alisma lanceolatum Althaea officinalis Anacamptis pyramidalis Anthriscus caucalis Aphanes arvensis* Apium graveolens Apium inundatum* Apium nodiflorum* Apium repens Aquilegia vulgaris* (3) Arum maculatum Asplenium obovatum* Aster linosyris Atriplex laciniata* Atriplex pedunculata Atriplex portulacoides Baldellia ranunculoides Berberis vulgaris§ Beta vulgaris Blackstonia perfoliata* Brassica nigra§* Bromus commutatus* Bromus racemosus Bryonia dioica* Bupleurum tenuissimum Callitriche obtusangula* Calystegia soldanella* Carduus nutans Carduus tenuiflorus* Carex davalliana Carex depauperata* Carex divisa* Carex divulsa Carex laevigata* Carex pendula* Carex punctata Carex strigosa Carex trinervis Carpinus betulus Carum verticillatum* Catapodium marinum* Catapodium rigidum* Cephalanthera damasonium Cerastium glomeratum (3) Ceratophyllum submersum Ceterach officinarum Chaerophyllum temulum* Chamaemelum nobile* Chenopodium chenopodioides Cicendia filiformis Cirsium acaule Cirsium dissectum* CDS PRESTON Cochlearia anglica Cochlearia pyrenaica* Coincya monensis* Corynephorus canescens Crataegus laevigata Crepis biennis Crepis capillaris (3) Crepis mollis Crithmum maritimum* Cruciata laevipes* Cuscuta epithymum (3) Cyperus fuscus Cytisus scoparius* (3) Dactylorhiza praetermissa Dianthus armeria Dipsacus fullonums* Dipsacus pilosus Dryopteris aemula* Dryopteris oreades* Dryopteris remota* Dryopteris submontana* Elytrigia atherica Epilobium hirsutum Epipactis leptochila* Epipactis phyllanthes Epipactis purpurata Equisetum telmateia* Erica mackaiana* Erodium lebelii* Euonymus europaeus Euphorbia hyberna* Euphorbia paralias* Euphorbia portlandica* Euphrasia tetraquetra* Festuca filiformis Festuca lemanii§* Filago lutescens§ Filago minima Filago vulgaris* Fritillaria meleagris§ Fumaria bastardii* Fumaria capreolata* Genista anglica* Genista pilosa Genista tinctoria Glaucium flavum Groenlandia densa Helianthemum oelandicum Homogyne alpina§* Hordeum marinum* Hordeum secalinum Hornungia petraea Hyacinthoides non-scripta* Hydrilla verticillata Hymenophyllum tunbrigense* Hypericum androsaemum* PLANTS WITH RANGE EDGES IN THE BRITISH ISLES Hypericum elodes* Hypericum humifusum Hypericum tetrapterum* Hypochaeris glabra Ilecebrum verticillatum* Inula conyzae* Isolepis cernua* Juncus foliosus* Juncus inflexus* Juncus maritimus* Juncus pygmaeus Juncus subnodulosus Koeleria macrantha* Lactuca virosa* Lamiastrum galeobdolon Lavatera arborea* Leontodon saxatilis* Lepidium heterophyllum* Lepidium latifolium (3) Limonium humile Limonium vulgare Lotus glaber Lotus pedunculatus Luronium natans Lysimachia nummularia (3) Malva moschata* (3) Marrubium vulgare (3) Meconopsis cambrica* Medicago minima Meum athamanticum* Minuartia sedoides* Moenchia erecta* Myosotis alpestris* Myosotis stolonifera* Myosotis sylvatica (3) Narcissus pseudonarcissus* Oenanthe crocata* Oenanthe fistulosa Oenanthe fluviatilis Oenanthe lachenalii* Ononis spinosa Ophrys apifera* Orchis laxiflora Orchis morio Orchis purpurea Orchis ustulata Ornithopus perpusillus* Orobanche alba Orobanche artemisiae-campestris Orobanche caryophyllacea Orobanche elatior Orobanche hederae* Orobanche minor (3) Orobanche purpurea Orobanche rapum-genistae* Orobanche reticulata Oxytropis halleri* Parapholis incurva* Parapholis strigosa Parentucellia viscosa* Parietaria judaica* Petasites hybridus (3) Phleum arenarium Phyteuma orbiculare Pinguicula lusitanica* Poa bulbosa Polypodium cambricum* Polypodium interjectum Polystichum setiferum* Potamogeton acutifolius Potamogeton coloratus Potamogeton trichoides Potentilla rupestris Potentilla sterilis* Primula elatior Puccinellia rupestris* Pulicaria dysenterica* Pulicaria vulgaris Ranunculus fluitans* Ranunculus omiophyllus* Ranunculus ophioglossifolius Ranunculus sardous§ Raphanus raphanistrum* Reseda lutea§* Rorippa microphylla * Rorippa nasturtium-aquaticum * Rosa agrestis Rosa arvensis* Rosa micrantha Rosa rubiginosa Rosa tomentosa Rumex conglomeratus* Rumex palustris Sagina apetala* Salix purpurea (3) Salvia pratensis§ Salvia verbenaca* Sanguisorba minor (3) Sarcocornia perennis* Saxifraga spathularis* Scabiosa columbaria Scrophularia auriculata* Scrophularia umbrosa Scutellaria minor* Seriphidium maritimum Sherardia arvensis (3) Silaum silaus Silene conica* Silene otites Sonchus palustris Sorbus torminalis Spergularia rupicola* 267 268 Spiranthes spiralis Stellaria neglecta* Stellaria pallida Symphytum tuberosum§* Teesdalia nudicaulis Teucrium scordium Teucrium scorodonia* Thlaspi caerulescens* Tilia platyphyllos Torilis nodosa* Trichomanes speciosum* Trifolium campestre Trifolium micranthum Trifolium scabrum* Trifolium striatum Trisetum flavescens (3) Ulex europaeus* Ulex gallii* Ulmus minor Umbilicus rupestris* Valeriana dioica Verbascum lychnitis Veronica catenata Veronica montana* Vicia lutea* Viola lutea* C. D. PRESTON Vulpia bromoides (3) Vulpia fasciculata* Wahlenbergia hederacea* Wolffia arrhiza SPECIES WITH NORTHERN EUROPEAN LIMITS BETWEEN 60° N AND 65° N AND NORTHERNMOST OCCURRENCES IN THE BRITISH ISLES These species reach their northern European limits north of 60° in Shetland, although Aphanes australis is only doubtfully native there (Scott & Palmer 1987). There are 257 other native British species that reach their northern limit in the 60—65° latitudinal band elsewhere in Europe. Aira caryophyllea* Aphanes australis* Deschampsia setacea* Eleogiton fluitans* Eryngium maritimum* Glyceria declinata* Isolepis setacea* Radiola linoides* Ranunculus hederaceus* Schoenus nigricans* APPENDIX 2. SPECIES AT OR NEAR THE SOUTHERN EDGE OF THEIR NATIVE EUROPEAN RANGE IN THE BRITISH ISLES Native British species which have their southern European limits in the latitudinal bands 55—60° N 50-55° N and 45—50° N are listed in this section. Species are. asterisked if the absolute limit is reached in the British Isles, but not if it they reach their southern limit at the same latitude in our area and in other European countries. § Doubtfully native (3) Limit(s) of native range particularly confused by itrodNetione (this also applies to doubtfully native taxa) SPECIES WITH SOUTHERN EUROPEAN LIMITS BETWEEN 55° N AND 60° N Artemisia norvegica* Carex rariflora* Carex salina* Carex saxatilis* Diapensia lapponica* Erigeron borealis* Euphrasia foulaensis* Koenigia islandica* Luzula arcuata* Minuartia rubella* Sagina nivalis* Salix arbuscula* Salix lanata* Salix myrsinites* Saxifraga rivularis* SPECIES WITH SOUTHERN EUROPEAN LIMITS BETWEEN 50° N AND 55° N Alchemilla wichurae Alopecurus borealis* Arenaria norvegica* Atriplex longipes* Atriplex praecox* Blysmus rufus Carex aquatilis* Cerastium arcticum* Cornus suecica Dactylorhiza purpurella* Euphrasia frigida* Euphrasia ostenfeldii* Euphrasia scottica* Festuca vivipara* Galium sterneri* PLANTS WITH RANGE EDGES IN THE BRITISH ISLES Hydrilla verticillata Juncus balticus Ligusticum scoticum* Mertensia maritima* Polygonum boreale* Rubus arcticus Salix phylicifolia* Saxifraga cespitosa* Saxifraga hypnoides* Saxifraga nivalis Sorbus rupicola* SPECIES WITH SOUTHERN EUROPEAN LIMITS BETWEEN 45° N AND 50° N As explained in the text, these species are not necessarily close to their southern limit in the British Isles, but as the list is so short it is included for completeness. Arabis petraea Atriplex glabriuscula Calamagrostis purpurea Calamagrostis stricta Callitriche hermaphroditica Carex magellanica Carex maritima Carex norvegica Carex pauciflora Centaurium littorale Cochlearia officinalis Crassula aquatica Dactylorhiza praetermissa Draba norvegica Euphrasia arctica Euphrasia confusa* Euphrasia tetraquetra Festuca armoricana Festuca huonii Festuca longifolia Gentianella uliginosa Juncus biglumis Juncus castaneus Leymus arenarius Limonium humile Limonium normannicum Luzula pallidula Minuartia stricta Najas flexilis Oenanthe fluviatilis Poa humilis Potamogeton rutilus Ribes spicatum Rubus chamaemorus Salicornia pusilla Saxifraga hirculus Saxifraga rosacea Tephroseris palustris Thymus serpyllum 269 ] | | I II i ; Aa is iit. Eg a ee Watsonia 26: 271—290 (2007) ‘Far from any house’ — assessing the status of doubtfully native species in the flora of the British Isles D. A. PEARMAN Algiers, Feock, Truro, Cornwall, TR3 6RA ABSTRACT The question of the native or alien status of British and Irish plants was a popular subject in Victorian times, but little serious work has been undertaken for a century; instead species have been looked at individually. A recent paper has dealt with archaeophytes; this deals with neophytes, those alien plants arriving after 1500. The advances in detailed distribution mapping (by grid squares) both in the British Isles and in much of N. Europe, the new research available in the fields of archaeobotany and in the interpretation of the nomenclature of early gardening works, both in medieval times and up to 1800, now allow a broader view to be taken. This can often be coupled with recent European work on alien and native status and on archaeophytes. It is suggested that a range of criteria are examined for each species, and although it is unlikely that any single one will give an answer, if several point in the same direction then a decision based on _ the probability is possible. The ten criteria used are: first date into cultivation; first date found in the wild; presence in semi-natural habitats; spatial distribution in Britain; trends in frequency; persistence; use; European range; archaeological evidence; genetic diversity. A range of species, all except one treated as alien or native or alien in the New Atlas of the British & Irish Flora, is examined illustrating the use of those criteria. Conservation considerations are discussed. KEYWORDS: Alien species; neophytes; European work; archaeobotany; medieval gardening; criteria; conservation considerations. INTRODUCTION The question of the status of British plants, whether a species is native or alien in the country as a whole, or whether it is native in one area but not in another, is bedevilled with uncertainty and optimism, especially the latter. The position in Ireland is usually, but not always, clearer. A recurring theme in very many County Floras is the sentiment that distance from habitation over-rules all other considerations, including common = sense. Whilst we were assembling the data from the recent New Atlas of the British & Irish Flora (Preston, Pearman & Dines 2002) it became apparent that for many species there was considerable divergence over views on their nativeness, even within the last 50 years, and that there had been no rigorous investigation, covering the whole of the flora, for well over a century, let alone any attempt to apply the same criteria to all the species in question. We therefore prepared a list of about 300 species (about 20% of the possibly native flora) where we felt there were questions to be raised over their origin, omitting many species whose status as aliens has never been questioned. We were inspired by the seminal paper by Webb (1985), where, briefly, he discussed eight criteria that could be used to produce a more objective assessment of native status than that currently used, and where he _ suggested examples of native species possibly requiring reconsideration. We subsequently used all those criteria except that on reproductive pattern, together with three others that we considered relevant. We analysed our list of 300 against the assignment of “nativeness” in the post-war standard floras and checklists, none of whom gave other than the most cursory reasons for their decisions. In fact there has been no comprehensive study work since the earlier work on the origins of plants by Watson (1832, 1847-1859, 1870) and Dunn (1905), and none at all that takes into account recent develop- ments that might be relevant in the world of horticulture and archaeology, or even of advances in mapping in Europe other than on a very few individual species. Our work was stimulated by the need to provide sound assessments for a complication which researchers did not have to face until comparatively recently, namely the fixation of the conservation industry with “nativeness”; that is the unwillingness to conserve something that might be “alien”, and therefore the pressure on commentators to label as native those plants that conservationists want to conserve or for which they have established S.S.9.1s. Di, D. A. PEARMAN We felt then that whilst it 1s difficult or even impossible to attain certainty in the field of origin, the issues were best served by setting out as many of the facts and dealing with as many of the facets as we could, and making a judgement based on all of those, and in repeating this same process, using the same criteria, for all the species in question. Obviously some criteria might be more important than others, not only as a generality, but for particular species, but that would only become clear in discussion. On this basis about 150 of these 300 species qualified as archaeophytes (that is plants which were brought to Britain by man, intentionally or unintentionally, and became naturalized there between the start of the Neolithic period [c.4000 BC], and AD 1500), and these are dealt with in Preston, Pearman & Hall (2004), where we set out instances from as many archaeo- botanical, horticultural and phytogeographical sources as we could. Of the remaining 150 we decided, again using as many criteria as possible, I. that 74 were neophytes (that is, species brought by man after 1500) or almost certainly so II. that another 44 were probably neophytes, though there were enough uncertainties to categorize them as either native or alien, and Ill. that the rest, though we had doubts over many, were left as natives in the hope that further work would assist in forming a more definite view. We are pleased that the conservation agencies have decided (Cheffings & Farrell 2005) that all the archaeophytes and all the species that we categorised as native or alien are to be treated as “honorary” natives for conservation purposes. In this paper I attempt to show the criteria that we used in coming to these decisions, supplemented by other sources which we have become aware of since 2000, especially from studies in Europe on alien species (based on ecological work to predict invasiveness) which introduce views on status there which are very often at odds with those given in standard floras. PREVIOUS TREATMENTS OF STATUS IN BRITAIN The knowledge of the total number of species in the British and Irish native flora grew steadily from the 16th century and was almost ~ complete (over 90%, excluding critical species) by the mid 19th century. Interest in the origins of this “native” flora arose in a more piecemeal way, and is well covered in Preston (2002). There was certainly understanding of the difference between the wild and the cultivated flora in the works of Turner and some, but rather less, in Gerard, Goodyer, Tradescent, Parkinson and Ray, up to 1700, and in Martyn, Withering and Hudson in the eighteenth century. These authors reported a plant was in the wild with only rare comments on whether they thought it might be other than a native plant, together with some cases of deliberate or accidental introductions. But Smith (1800-1804, 1828), both in his own works and in conjunction with Sowerby in their illustrated English Botany (Smith & Sowerby 1790-1814), complicated the study of nativeness by admitting into those works (possibly because they were suitable subjects for illustration) a host of species that were little more than garden escapes and even deliberate plantings or errors. Despite not putting forward any rigorous evidence he had a considerable and lasting influence on his contemporaries. For instance the distinguished Henry Aiton, in his first edition of the plants cultivated at Kew (Aiton 1789), gave the origin of many plants we now call alien as from overseas, with details of who had introduced them and often the date of this. But by the second edition (Aiton 1810-1813), that is after Smith’s first works, he had altered the country of origin to “Britain”, and this was followed by the all the subsequent enumerators of plants grown in Britain (Loudon, Sweet, Paxton ef al.). In Druce’s paper on the Extinct and Dubious Plants of Britain (1919) over 50 of the 147 species covered had their origin in Smith & Sowerby. An example of the text in Smith & Sowerby is too apposite to omit in this paper. Having discussed Leucojum aestivum, which a lady from Suffolk described as a troublesome weed in old pastures that had never been broken up, he passes to Fritillary, where he writes “If we can allow the Leucojum a place in a work on British plants we cannot hesitate about the Fritillaria, which although not noticed by Ray or Dillenius, is very common in parts of Middlesex, as well as Suffolk and other counties. Mrs Cobbald sent it from Little Stoneham (Suffolk) with Leucojum aestivum, Ornithogalum umbellatum and Colchicum”. None of these is in a native site, other than possibly the Colchicum! Again and again the STATUS OF DOUBTFULLY NATIVE SPECIES 23 species in question is described as “far from any house”, “not the slightest doubt of its being indigenous” or “the situation seems to preclude the idea of an escape from gardens” and the like. It was not until Watson, in the second quarter of the nineteenth century, that there was any serious attempt to stand back and consider whether there was any evidence on the likely origin of particular species in the British flora. Only four years after the publication of Smith’s English Flora, Watson began this process of more critically assessing the origin of species, in his Outlines of the Geographical Distrib- ution of British Plants (Watson 1832), a process elaborated and refined in his major work, Cybele Britannica (Watson 1847-1859). Here he divided alien species into three categories, using the following names for them: Alien: “now more or less established, but either presumed or certainly known to have been originally introduced from _ other countries”’. Denizen: “at present maintaining its habitats, as if a native, without the aid of man, yet liable to some suspicion of having been originally introduced”. Colonist: “a weed of cultivated land or about houses, and seldom found except in places Where the ground has been adapted for its production by the operations of man”. In his Compendium (Watson 1870) he later added a fourth category, such alien place or Casual, for “chance stragglers .... species aS are uncertain in persistence”’. The subject, and its principle relevance to the treatment of archaeophytes, is discussed in more detail in Preston et al. (2004). His treatment of neophytes was much simpler. Of the 74 species selected by us from our list as neophytes, Watson covered 48, and of these he categorised only four as native without any caveats. Of the 44 species we selected as native or alien, he covered 34, and called 17 native with no caveats. Watson’s work was followed in virtually every county flora and national check-list for the next 100 years, including Druce’s check- lists (1908, 1928) and his Comital Flora (1932). Certainly there were local exceptions and special pleadings, but the general framework was either accepted or deferred to. The problem, which affects too many local flora writers, is the lack of experience of the flora of Britain and Ireland as a whole, a problem we found still endemic when preparing the New Atlas (Preston et al. 2002) where adjacent counties assigned different statuses to the same species. Watson, and later Druce, by looking at the flora as a whole, was able to see the inconsistencies and comment on them. Another work of that period was that of Dunn (1905) on the alien flora of Britain, where he covered in short accounts all the species that he considered alien. He dealt with 924 species, together with a further 170 that he had some doubts over, but accepted as probably native. In his book he described the habitat, mentioned the first date in the wild if he thought it relevant and discussed the wider distribution. His introduction covers many of the points covered in Preston ef al. (2004) and is still relevant. He suggests that “it is a general experience, and one that is to be expected, that two areas inhabited by a given native species are seldom known to be separated by a large tract of similar, and apparently suitable ground, devoid of that species”. He qualifies this by agreeing that it was not impossible that such a gap might occur in the truly native range, for instance as a consequence of approaching extinction, but recounts the study of geo- graphical distribution to shed light on these conundrums — an approach we tried to bring together in the New Atlas (Preston et al. 2002). Dunn’s work, other than in the context of the general acceptance of Watson’s works, receiv- ed little attention at the time, though advances in the studies covered by my criteria have largely supported his conclusions. Watson’s position held sway until after World War II and Lousley (1953), in a review of the classification of alien plants, espoused his main points with only minor modifications. However, for reasons I cannot really pin down, Watson’s system was falling out of favour by then, though a few local floras after 1960 have used it. It would be difficult to say that Watson’s tenets had been replaced by fresh reasoning in the post-war British floras and check-lists, because there is no discussion in any of them - of the reasons behind their categorization of statuses, which are limited to “native” or “introduced” with qualifiers. In the Flora of the British Isles (Clapham et al. 1952) there is only a reference in the glossary, where “introduced” is defined (in the appendix, p. 1521) as plants 274 D. A. PEARMAN “known to have been, or strongly suspected of having been, brought into the British Is. accidentally or intentionally by man within historic times”. Later editions (Clapham ef al. 1962, 1987) modify the status of some of the treatments of individual species, but still shed no light on the reasons. Dandy (1958) gives no explanations, merely mentioning the difficulty of differentiating native and long-established aliens. I cannot find a rationale or explanation in Stace (1991, 1997), though I am fully aware that he not only has a huge personal experience but also many active correspondents. Kent (1992) gives no explanations at all; Clement & Foster (1994) and Ryves, Clement & Foster (1996) treat ancient introductions as native and for other species either deal with them as aliens, or, if in doubt, square bracket the species with the comment “accepted, with reservations, as native”. So, whilst it is possible to see there, for instance, that Pentaglottis sempervirens 1s treated as a native or an ancient introduction, it is not possible to see why. I have also consulted Reynolds (2002) for the treatment of Ireland. Every plant on our draft list of neophytes and of native or alien was treated by her as alien in Ireland other than three instances where, in following the Irish Census Catalogue, the status in Ireland was not further questioned by her. It is not altogether surprising that the above dealt so summarily with aliens or doubtful aliens as their works were all fulfilling many other purposes and status was only one, and a mixed one at that. But the real point is that I cannot trace any attempt since Watson and Dunn to examine a range of species, using as many criteria as possible, and to present that information, and this is the purpose of this paper. CRITERIA USED AND THEIR RELEVANCE 1. FIRST DATE IN CULTIVATION The main sources of summarised information are Aiton (1810-1813) and Loudon (1855). The second is later but omits the invaluable details on sources contained in the former. Elucidation of early sources (Turner and his predecessors, Gerard ef al.) usually but not always contained in Aiton and Loudon, has been made much clearer by the recent works of John Harvey and his colleagues, covering both interpretation of nomenclature and previously unknown or _ inaccessible works. More background is contained in the introductory chapter in Preston et al. (2002). Work on species introduced since Loudon has been hampered by the fact that nothing comprehensive has been attempted since his last edition, possibly because there were so many, so information has had to have been assembled from a great variety of sources. The other most significant missing link in our researches is the history of grass seed and fodder crop introductions, together with the impurities that these might have brought with them (see Rich & Karran 2003) for a rare illustration). I consider that a date in cultivation for a species that is considerably earlier than the first date for the wild is very relevant for garden species. (e.g. Aconitum napellus, Allium sphaerocephalum, Daphne mezereum, Fritillaria meleagris, Galanthus nivalis) but, of course, is less relevant for a species that might have been grown for curiosity but never entered mainstream gardening (e.g. Linaria supina, Hypericum canadense, Serapias lingua). Note, as discussed earlier, that once a species was shown, particularly by Smith & Sowerby, as native in Britain, then Aiton, Loudon and others no longer showed its source and introduction date. 2. FIRST DATE IN THE WILD The information to support the first dates of discovery comes principally from Clarke (1900), Dunn (1905) and Druce (1932), and as we said in the New Atlas, we regard the first two as more reliable than the last. More background is contained in the introductory chapter in Preston et al. (2002). As with work on introductions into cultivation, so any research on first dates in the wild has similarly lapsed, with virtually no work taking place in the last 70 years, despite the increased interest in alien plants generally. Since the New Atlas further work has seen the emendation of about 15% of the dates given there, but little further herbarium work, which is potentially the main remaining source of better data, has been undertaken. For species other than those that are critical or really insignificant, I consider the first date in the wild as of very great importance, in helping to assess possible arrival dates, particularly in well-worked areas of British Isles. Of the 74 species in our initial selection that we decided were probably neophytes, 44 had first dates in the wild in the 19th and 20th centuries, with a further 19 in the 18th century, mostly after 1750. STATUS OF DOUBTFULLY NATIVE SPECIES Dil 3. PRESENCE IN SEMI-NATURAL HABITATS AND IN PLANT COMMUNITIES We collected information for these for the paper on archaeophytes (Preston et al. 2004), but because the species covered by this paper are such an eclectic mix compared with those we selected as archaeophytes their occurrence or not in semi-natural habitats is of less relevance, and probably of more importance for the non-garden and annual plants. Nonetheless, there must be a reasonable hypothesis to suggest how species have survived from their arrival in the British Isles (usually earlier in post-glacial times) and this is often very difficult for species in man-made _ habitats. Thus if a plant grows only in man-made habitats it is likely to be alien. In each case Rodwell (1991-2000) has been consulted for the habitats of species, and has been supplemented by the information in Hill, Preston & Roy (2004), from county floras and by personal knowledge. It would be no surprise that many of the species illustrated in the following case studies did not appear in Rodwell, as almost all are rare. 4. SPATIAL COHERENCE IN BRITAIN AND IRELAND This is an interpretation of the actual range in Britain and Ireland, compared with the available habitat. Thus Stachys alpina, restricted to two copses when it could occur anywhere, has no coherence whatsoever, and Galanthus nivalis, omnipresent, has. In itself, of course, a limited range, when there is plenty of other suitable habitat, is not conclusive to a decision on alien status or not, but it is indicative. Where a species, such as Euphorbia serrulata, has a core “native” range and other more scattered records elsewhere, only that alleged “native” range is taken into account. 5. TRENDS IN FREQUENCY (DYNAMISM) In our original selection we looked also at dynamism in Britain (whether a species was increasing or decreasing, and if so, how fast). Rapid increase is frequently one of the hallmarks of a neophyte, whereas natives and archaeophytes, in general, show slow declines. Furthermore if the environment changes, then the range of natives tend to change too, whereas the range of aliens is changing even if the environment stays the same. A further subject for research on which some progress has been made in Europe, but as yet little here, is the time-lag between the arrival of an alien plant in cultivation, and its appearance, Nn if it makes that transition, in the wild. Recent work in Germany (Kowarik 1995) (only one of a number of recent papers) gives a surprisingly long time-lag for woody plants, so we might only now being seeing the rapid spread of plants introduced 100 to 200 years ago. 6. PERSISTENCE This covers whether the recorded sites were long-lasting or ephemeral. As a generalisation many aliens are ephemeral, their presence characterised by a number of different sites, each short-lived. 7. USE This covers whether the species was grown in gardens, as an herbal or medicinal plant, or as a crop, and thus whether it is more likely that it has escaped into the wild. Of course a native plant may be used by man — there are very many medical and culinary examples — but if there is a use, then there is a better chance of it becoming established in the wild, and, again, this is just one of a range of criteria. 8. EUROPEAN RANGE Most of our standard floras give distribution in Europe, or in the rest of the world, or both. The information will have been obtained from foreign national floras, and from regional and local floras, and more recently atlases such as Meusel et al. (1965, 1978, 1992) and Hultén & Fries (1986), Bolos & Vigo (1984-2001), and others where the distribution maps have the range shaded could have been used. Whilst this last enables the total range to be seen, it often fails to differentiate between the core range and outliers. Webb (1965) recognised this as a problem, writing that “the elegant loops and curves which we see in so many maps conceal ambiguities and ignorance”! However since Perring & Walters (1962), and the earlier work by Hultén (1950) a few countries have started to produce grid-based atlases which enable the real distribution to be seen much more clearly, and to better allow us (or more easily than a text can) to evaluate our flora with its distribution elsewhere. As my survey of these works is new and perforce incomplete, in that either I am still unaware of the existence of data, or, more likely, the data that I consider necessary is not available, this must be considered work-in- progress, but it is of such major assistance to this paper that it will surely be used more and more. 276 D. A. PEARMAN There are recent grid-based Atlases, all at a scale of 10 x 10 km or larger, for Germany (one each for the former West and East) (Haupler & Schonfelder 1988; Benkert ef al. 1996), for Holland (Mennema ef al. 1980, 1985; Meiden et al. 1989) and an incomplete series (covering about 900 species) for N.E. France, covering an area from Cherbourg in Normandy to the German border, down to Paris, and including Luxemburg, most of Belgium and much of Holland (institut Floristique Franco-Belge 1978-2001). There is also available a volume of maps for the Breton department of Finistaire (Hardy 2002), useful for Atlantic species. Atlas Florae Europaeae (Jalas et al. 1972- 1994, 1996, 1999) covers all species in Vol. 1 in Flora Europaea_ (roughly, up_ to Saxifragaceae) and the latest (Kurtto ef al. 2004) volume starts the Rosaceae. These are, of course, invaluable, but since they are at the scale of 50 x 50 km, show less detail than the country atlases listed above. A further source of research, which is sometimes not available from the sources cited above, is information on how European national and regional floras describe the status of our flora in their countries. Often of course, their statements have been as vacillatory as ours, and though, again, I feel that I have barely explored this subject, I have been able to consult interesting lists for countries which, like ours, could well be at the edge of the range of Mediterranean species. In particular, I have used the recent checklists for Germany (FloraWeb — http:/ice.zadi.de/floraweb), for Poland (Mirek et al. 1995) and the Czech Republic (PySek et al. 2002). This whole subject, linked to work exploring and predict- ing the ecology of plant invasions, is developing apace, and Pysak ef al. (2002) make the interesting point of the “remarkable difference they found between the data drawn from the standard floras and (that from) checklists commenting on species immigration status and studies focussing specifically on alien plants”. In other words a study of work in other countries is often essential to a better understanding of the origin of our own flora, in the same way as a national Atlas might see a broader picture than a county Flora. 9, ARCHAEOLOGICAL EVIDENCE Of major importance in the paper on archaeophytes (Preston et al. 2004) was the information available on archaeological evidence, almost all of it gathered in the last 30 years. For neophytes the evidence is, not surprisingly, almost entirely absent. Of all the species investigated for this paper the only evidence found was for the currants, Ribes nigrum and R. uva-crispa, where post-Norman remains were identified. 10. GENETICS There has been relatively little relevant work here. Nevertheless there has been research on Draba aizoides (John 1992) which seemed to support its claim to nativeness and for Leucojum aestivum where David Coombe was able do demonstrate differences between the native and alien subspecies (FitzGerald, in Gillam 1993). CASE STUDIES For our work for the New Atlas (Preston et al. 2002) we looked at all the criteria for each species before coming to a decision, using a “score” by which we decided the probable status of each. A spreadsheet, showing all the species we covered in the categories that I have described (neophyte, native or alien and others) and using the criteria outlined above, is available from the author. Rather than produce that very large table it seems more appropriate to try and give a selection of examples to illustrate the use of the criteria. Each study is prefaced by a heading and condensed paragraph setting out the information available on the relevant criteria, though the information on the European distribution is dealt with in the main text that follows. In this paragraph “NVC” refers to the information in Rodwell (1991-2000); “Watson” to Watson (1847-1859, 1870), “Dunn” to Dunn (1905), “Stace” to Stace (1991), “Kent”? to Kent (1992), @lementrc Foster” to Clement & Foster (1994), “Fl Europaea” to Tutin et al. (1968-1980, 1993), and “VCCC” to Stace et al. (2003). A garden plant, long cultivated here, but with late first records, and uncertain native range EUPHORBIA CYPARISSIAS In cultivation by 1640, first record in wild 1796 (Earl of Stamford’s woods at Enville, Staffs). It is not included in the NVC and is invariably a species of open ground. The records are very scattered; it is increasing, and it is persistent in its sites. It is grown as a garden plant. It was STATUS OF DOUBTFULLY NATIVE SPECIES PATI treated by Watson as alien, and by Dunn as native. The treatment in recent floras has varied, with some showing as native, some alien, and F/ Europaea and the VCCC as alien. This species is frequent in continental Europe, up to, but not within 20 miles of, the Channel coast, so a first glance at a world distribution map would not arouse many doubts on nativeness, though Mennema et al. (1980) say it has been extending northwards for some centuries. Yet a combination of all the above criteria points decisively against this. Even Dunn’s decision is not what it appears. He says... “not, the only natural locality is on the downs near Dover” (Dunn 1905). Yet this is presumably based on the entry in the Flora of Kent (Hanbury & Marshall 1899) which actually says “perhaps native in one station.....chalky hill-slope near Dover. ..it may be indigenous here; the continental distribution of the species rather favours this view.” Druce (1908) considered it an alien and Clement & Foster (1994) report, inter alia, that it was formerly brought in with racehorses. It is a very persistent garden plant, spreading through its rhizomes on light soils. Decision: alien (neophyte). A garden plant, long cultivated here, but with late first records, and uncertain native range FRITILLARIA MELEAGRIS In cultivation at least by 1597, and possibly by iaiswawithe first record m wild - 1736 (Middlesex, Harefield). NVC MGé4, and widely naturalised elsewhere. Spatially it is fairly widespread, but in most sites it has declined though it can multiply quickly given appropriate conditions. A well-known garden plant, treated by Watson as a denizen and not covered in Dunn. All recent floras other than Kent and Stace (1997) have treated it as native, as has Flora Europaea and the VCCC. More has been written about the nativeness or otherwise of this species than any other. The case for was recently summarised in Oswald (1992) and that against by Harvey, the expert on medieval plants (1996), and I refer readers to those papers for the case in full (and in Harvey's case for much more besides — endlessly provoking and utterly fascinating). A summary for nativeness (Oswald) might be: 1. Near-continental distribution makes extension into southern England possible. 2. Late discovery possibly due to very short early flowering time and restricted range. 3. Habitat often waterlogged in winter and not the sort of countryside a 17th century botanist might have felt exploring. 4. A reference in the 17th century naturalist John Aubrey to a plant called ‘crow-bells’ might represent a very much earlier record. To these should be added, of course, the fact that many of the sites are first-rate examples of neutral grassland and that aliens are very rare in this habitat. Harvey’s points include: 1. Late date for such a colourful and obvious flower, and a dismissal of any connection with the plant described by Aubrey. 2. Unlikely to have been overlooked by Turner, Gerard, Parkinson, Johnson, Ray, Morison and Dillenius. Since the publication of those two papers more has come to light regarding the continental distribution. Preston et al. (2004) point out that it is treated as native in Poland, but an archaeophyte in Germany, a neophyte in Finland and a casual in the Czech Republic. In Demark, cited by Harvey, it is treated as a neophyte, arriving in 1647, and in Sweden (Zhang 1983) a neophyte, arriving in 1742 or 1743, having been grown in Uppsala Botanic Garden since at least 1658. Here it has spread into flower-rich meadows, behaving like a native if one did not know its source. Finally, it is absent from N.E. France (Institut Floristique Franco-Belge 1978-2001) and in Brittany is found only in the south (des Abbayes ef al. 1971). So the European distribution militates strongly against it being a native here. In England the first date in the wild is 1736, from a site where it had been known for “forty years”. We have always ignored these feats of memory as being unreliable, but the next wild record I can trace is not until 1776 (Hind 1889), where it was described as abundant in meadows. Then, for the 29 counties where it was mapped as native in the New Atlas (and some of those are very recent discoveries indeed), it was found in another twelve in the next 30 years, ranging from Stafford to Suffolk and down to Dorset, and another nine in the next 25 years. It was not discovered until 1862 in the county possibly most celebrated for its display, Wiltshire, and the Herefordshire and Huntindon “native” sites are both twentieth century finds. What happened either to account for collective myopia all over southern England, or 278 D. A. PEARMAN to explain why is was discovered in twelve out of 29 supposedly native counties within 30 years? I can offer no explanation. Its propensity to multiply very fast is well-recorded (e.g. Zhang 1983), and Kevin Walker (in litt) tells me of his experiences at Portholme in Hunts. At that site there was very little in the years 1925 to 1980, and then a vast increase in the period 1980 to date. It seemed to be helped by winter flooding (spreading the seeds) and by hay cuts (seeds dispersed). How could it arrive within a very few years other than by repeated introductions? Decision: alien (neophyte). A garden plant, long cultivated here, but with late first records, and uncertain native range RIBES UVA-CRISPA In cultivation by 1275, first record in wild 1763 (Cambs, though both Babington (1860) and Perring et al. (1964) considered it doubtfully native there). In NVC (W8 (Fraxinus-Acer campestre woods) but only as a very incidental constituent and elsewhere a species of varied habitats. The records are widespread, increasing (possibly as a result of better recording of aliens), and persistent. It has been grown for centuries and was treated by Watson as a denizen, and by Dunn as a native. All recent floras treat as native, with some doubts in Stace, but the VCCC treats as alien. There is archaeological evidence from medieval times (1225-1400) (Tomlinson & Hall 1996). Roach (1985) recorded a pale (.e. green) form arriving from Flanders in 1509. Turner (1548) has it growing only in gardens, though he mentions that in Germany it grows in the fields amongst other bushes. Gerard (1597) describes it as well-established (in gardens) but said it had no name amongst old writers who either knew it not or esteemed it not. Smith & Sowerby (1790-1814) say “nothing can be more difficult than to say whether this plant can be truly a native of Britain. It is so far naturalised as to be common everywhere. Mr Robson finds it plentifully in woods and hedges around Darlington, which may be its native country’. Dunn (1905) describes it as common in damp woods in perfectly natural habitats, though undoubtedly alien in many places. It seeds very easily and as with R. rubrum, it might always have existed in a wild and weedy form but if so it is odd that it was ignored by all the early botanists. Decision: alien (neophyte, though possible archaeophyte). A species once grown in gardens, and capable of being spread by seeds, long-established in one site seemingly as a component of semi- natural vegetation LONICERA XYLOSTEUM In cultivation 1596, first record in wild 1770 (Northumberland, but by 1801 in Sussex, nr Arundel). It is not included in the NVC, but occurs in ancient woodland and hedgerows at one site. Older records particularly were fairly widespread, but these were mainly casual records, and it is only locally persistent. It was formerly grown as a garden plant, but now rarely as more floriferous species have been introduced. Watson decided as probably alien, Dunn as a native. Recent floras and FI Europaea have been equally divided between native and alien status. It is widely distributed in Europe, principally in higher areas away from the coastal plains, as far north and west as just above Paris, and in Germany up to Hanover. Elsewhere, as in Brittany (des Abbayes et al.), it is as a casual or introduction. In Wigginton (1999) much is made of the habitat in species-rich woodland or hedgerows, often near to old trackways, and ancient woodland herbaceous associates are given. In particular the presence of stools of Tilia platyphyllos is stressed, and paralleled with similar woods in France. Webb (1985) singles this species out for special attention under his criteria of “frequency of known naturalization”. After pointing out that if a plant claimed to be native in one locality is becoming more and more widely naturalised in similar habitats elsewhere then some reconsideration is called for, he points out that the claim for nativeness in one station near Arundel “can be traced back to a statement by Borrer, published by Smith in 1801, that it was “growing plentifully and certainly wild’. But Borrer was only 19 at the time.....Smith admitted at the time that it was frequent in gardens and that he had previously regarded it as an escape........ I cannot believe that the statement of the youthful Borrer, copied uncritically from one book to another, outweighs the mass of evidence in the opposite direction”. It is still in the Arundel site, but the evidence (late discovery, use as garden plant and freq- uency of introduction elsewhere) sides with STATUS OF DOUBTFULLY NATIVE SPECIES 279 Webb (and Watson before him, whom Webb used but did not acknowledge!). Decision: alien (neophyte). A well-known garden plant, with a very late discovery date, with an allegedly native endemic subspecies ACONITUM NAPELLUS In cultivation by 1596, first record in wild 1819 (Herefords, “in a truly wild state”). It is not included in the NVC but often forms an integral part of riparian vegetation. Spatially it is limited as a presumed native and _ its dynamism is confused by escapes; it is very persistent. It has been grown as a garden plant for centuries. Watson treated it as a probable denizen and Dunn as an alien. Recent floras all treat it as a native, possibly influenced by the suggestion of an endemic subspecies. The taxonomy of this species is complicated. Jalas & Suominen (1972-1994) follow one taxonomist’s recent work in showing the British species as a near endemic subspecies, subsp. napellus, restricted to western Britain with one outlier in S.W. France. Stace (1997) seems ambivalent. The Brittany flora (des Abbayes et al. 1971) shows all records there as adventive. It is clear there is no consensus and that more work is needed to establish whether there actually is any endemic subspecies here. I have traced no map of the European range, other than a shaded map in Bolos & Vigo (1984-2001) which shows the _ species occurring from central and southern France across to Switzerland and northern Spain. It was grown in Elizabethan times, and it seems inconceivable that, as a native, it was not discovered in the wild until 1819. It has many strong advocates as a native — there is a notable passage in the excellent Flora of Bristol (White 1912). It has the appearance of being perfectly wild along shady streams, but all the sites, some long-standing, investigated by the author in Dorset and Somerset turn out to be below gardens, old or current, and pieces of the tuberous rhizomes, washed away by floods, readily establish on open ground downstream. Incidentally White was a strong optimist over plant status — he gave the benefit of doubt to almost every relevant species covered in this paper, including the following. The very late discovery, coupled with the large gap in the potential native range, seem to outweigh the semi-natural niche that it is now found. The taxonomy still needs to be resolved. Decision: alien (neophyte). Work needed to clarify the taxonomy. A species, once grown as a garden plant, with a very late date of discovery and a very odd distribution STACHYS ALPINA In cultivation by 1597, first record in wild 1897 (Glos, Wooton-under-Edge). It occurs in woodland and on verges. As a presumed native its distribution is extremely limited, and its dynamism is uncertain, as it is obscured by conservation action; it is persistent given favourable conditions. It has been used as a garden plant, but probably not now, and was not covered by Watson or Dunn. All recent floras describe it as native. The European distribution is interesting — primarily around the Alps, but with a band across central Germany, Belgium and mid France, with further outliers N.W. of Paris and in S Brittany, though the most recent Belgium flora (Lambinon et al. 2004) remarks it is sometimes introduced. It has been closely studied by Kay & John (1995), who report that all the populations are closely related and almost monomorphic, that all the sites have nothing out of the ordinary to explain such a refuge, that it sets seed easily and spreads by this and root division, and that following the discovery of the Gloucester site, where it considerably increased in range following discovery, many botanists grew it in their gardens. As such they feel that it is very unlikely indeed to be a native plant. Though it does seem to be a garden plant of very little value, it seems totally unlikely that its British distribution should be only three localities, in habitats that are either secondary woodland or ruderal sites. I am amazed its nativeness has not been questioned before! Decision: alien (neophyte). A species found in semi-natural grassland, with a coherent European distribution, but a late discovery PRUNELLA LACINIATA This was cultivated in 1713, but not found in the wild until 1886 (Glos). It is found in semi- natural grassland. It has been recorded in over. 60 10-km squares, mainly in a broad swathe of S England, but is declining. Watson did not cover it, but Dunn treated it as an alien, as have most recent floras, though Stace treats as possibly native. 280 D. A. PEARMAN Hultén & Fries (1986) show a wide swathe of distribution across Europe, reaching N.E. France and southern England, but in fact Institut Floristique Franco-Belge (1978-2001) shows only very scattered records away from Belgium. There are records of the species being introduced with clover seed (Preston ef al. 2002), and very many of the remaining sites in England have populations of the hybrid with P. vulgaris; 1n fact this seems to lead to the extinction of P. laciniata. (e.g. Morton 1973). This lack of a long-term survival mechanism, coupled with the classic curve of a frequent repeated introduction and almost terminal decline, together with the very late date of discovery, all points conclusively to an introduction. Decision: alien (neophyte). A late discovery with no contiguous European range and no real semi-natural habitat EUPHORBIA SERRULATA There are no historic details of cultivation, and first record in wild is 1773 (Tintern). It seems to be a species of open disturbed habitats, only growing in woodlands when there is little competition and plenty of light. Its area of presumed nativeness is very restricted indeed and it has littke dynamism, decreasing other than as a casual. It is usually persistent in open habitats. It is certainly grown as a garden plant, but possibly only recently. It was treated by Watson as a native in the Wye Valley, it is not included in Dunn, and all recent floras all treat it as native, other than Kent. Due to past confusion with E. platycarpos its history as a garden plant is not recorded. It is a weed species, responding very well to disturbance and light. In garden conditions, both in alkaline Dorset (pH 8-1) and acid Cornwall (pH 4:5) it seeds and _ spreads prolifically. But it is confined as a presumed native to around 20 historical sites in the Wye Valley, with an increasing number of escapes elsewhere. I have not found any map of the world distribution, but map floras of West Germany (Haupler & Schonfelder 1988) and N.E. France (Institut Floristique Franco-Belge 1978—2001)) show a central European distribution with no occurrences north the Cologne area in Germany and in France from Lorraine, where it is frequent, westwards, with very scattered records to southern Brittany (Mayenne). It is supposed to be a calcicole, and its very limited range, absence from anywhere else in Britain and northern Europe, plus its late discovery, justifies the doubt over status. Decision: alien (neophyte). A weedy species of open habitats, though with a Mediterranean-Atlantic distribution, and some persistence LAVATERA CRETICA In cultivation 1723, with first record in wild 1859 (Surrey, as a grain impurity, but 1873 in Scilly). It is a species of open disturbed ground. Its distribution shows very little spread, little dynamism, but it is persistent in Scilly and the Channel Isles. It has no use, is not covered in Watson, and is treated by Dunn as a casual. All recent floras, other than the VCCC, show it as native. Its European range is Mediterranean (Bolos & Vigo 1984-2001), reaching Portugal, with the French records possibly (the map is unclear) confined to the far S.W. and the far N.W. It is rare, sporadic and non-persistent in mainland Cornwall; in Scilly persistent (first discovered in 1873) and restricted to disturbed ground (“waste ground, old quarries, roadsides and bulbfields”’). In Jersey it was first recorded as a casual in 1879 (Le Sueur 1984), and was then rare over the next 80 years, and in Guernsey it was found before 1886 (McClintock 1975), and has spread only in the last 50 years. Lousley (1971) argues that failure to spread in Scilly indicates nativeness but where did it grow before the modern landscape was formed? He further claims that it occurs up the west coast of France in habits similar to those in Scilly, but this is not born out by the Brittany flora (des Abbayes et al. 1971) where it is described as a naturalised adventive or occasional casual. A common Mediterranean weed, but a most unlikely native plant. Decision: alien (neophyte). A weedy species of open habitats, though with a Mediterranean-Atlantic distribution ANISANTHA MADRITENSIS Not apparently cultivated and first record in wild 1716 (Kent, Sandown Castle, but known in Avon Gorge since 1773). It is a species of open disturbed ground. Fairly widespread, with some dynamism but these are mainly casual records and it is persistent only in two core STATUS OF DOUBTFULLY NATIVE SPECIES 281 areas. Use, none, and treated by Watson as possibly native, with caveats, but by Dunn as an alien. Recent floras are completely contradictory. A frequent weed in the Mediterranean, and occasional up the west coast of France to southern Brittany; absent from Germany and a casual elsewhere in northern Europe. Perring & Walters (1962) claim “native or long established” in Pembrokeshire and Glamorgan, together with the Avon Gorge, but Ellis (1983) regards it as colonist from England, with all the older records from there being from around ports or ancient buildings. The case for the Avon Gorge is certainly more convincing but is more likely to be an early introduction in view of the European distribution, the commercial importance of area and subsequent spread, a view endorsed by Lovatt (1982). Decision: alien (neophyte). A weedy species of open habitats SPERGULARIA BOCCONEI There is no record of cultivation; first record in wild 1901 (Cornwall, Par). It is a species of open disturbed ground. Spatially very limited as a presumed native, with little dynamism, declining and not persistent. Use, none, and not covered by Watson or by Dunn. Recent floras have treated it as native, with some doubts, notably in Stace, and the VCCC shows it as alien. A predominately coastal species, found around the Mediterranean (and eastwards to Iran), and very rare in France (Jalas & Suominen 1972-1994). It is found in the Channel Isles (Guernsey 1912 and Jersey 1906), where it is considered a _ probable introduction (McClintock 1975; Le Sueur 1984). Its other British sites are mostly at or near ports, where its usual habitat seems to be car parks and in other weedy places; few (any?) of the populations are long-persistent. Decision: alien (neophyte). A weedy species of open habitats, with a coherent European distribution PETRORHAGIA PROLIFERA In cultivation, no record, first record in wild not known for sure due to confusion with P. nanteuilii, but at least by 1840 (Norfolk). It is a species of open disturbed ground. It is only currently recorded from two sites, is declining, though it has been known in its current areas for some time. There is no recorded use, and it was not covered by Watson or Dunn. Of recent floras only Kent calls it native, the rest equivocate. This species has a coherent European distribution (Jalas & Suominen 1972-1994), reaching the Channel coast, though Lambinon et al. (2004) describe it as only in southern Belgium and Lorraine, often introduced elsewhere. Mennema et al. (1980) show its habitat in Holland as in dry open sandy places along the great rivers. It is a weed in the southern Hemisphere. In the area of Norfolk where one of its two British sites occur, it has been known for 160 years, the remaining site is next to a concrete roadway, and, although it is in the adjacent semi-natural grassland, Petch & Swann (1968) described it as a casual there. In its other site in Bedfordshire it is by a sand pit, adjacent to a railway. Akeroyd & Beckett (1995) argue for its native status at these sites, but its late date of discovery (even allowing for confusion with P. nanteuilii), its single site with any claim to continuity (out of only two that have even been suggested as native) and its habitat, all have greater weight than the continental distribution. Decision: alien (neophyte). A weedy species of open habitats, long-known in Britain, but probably never more than a casual other than in the last century at one site TORDYLIUM MAXIMUM There are no records of cultivation but the first record in wild is 1670 (Middlesex, but first in Essex 1875). It is a species of open disturbed ground. Spatially very rare and scattered and declining. It has some persistence, but this is now aided and obscured by conservation intervention. It has no uses, and Watson treated it as alien or denizen and Dunn as alien. All recent floras, including Fl Europaea treat it as alien except Clement & Foster, possibly native, and the VCCC, which treats it as native. Its European range is Mediterranean, reaching across France to Brittany, but not in N.E. France, Belgium or Holland (Bolos & Vigo (1984-2001). Adams (1999) seems incorrect in stating it is considered native in northern France. Institut Floristique Franco- Belge (1978-2001) show it as a rare casual, with only a very few records, all to the south of Paris. It is also a casual in Holland and Germany. Smith & Sowerby (1790-1814) give only an old record near Oxford, adding “but having been observed there for above a century, it cannot but be considered as a native’. Adams 282 (1999) postulates that it might be an overlooked native that firmly established itself during (or perhaps before) the climatic optimum of the sixteenth century, and points to its presence with other continental therm- ophiles such as Lactuca saligna and Vicia bithynica. However, in Britain this species has always been a rare casual and nothing more, other than in Essex where it has persisted in a number of open sites adjacent to forts and other disturbed areas, though the true position is hopelessly confused by planting and gardening. Decision: alien (neophyte). A weedy species with coherent European distribution, but late discovery and habitat suggest a recent arrival CERASTIUM BRACHYPETALUM There is no record of cultivation, with the first record in the wild 1947 (Beds). Abroad, it is a species of open disturbed ground. Its distribution is extremely limited as a presumed native, with no dynamism at all, and persistent only with conservation action. It was not covered by Watson or Dunn. Some recent floras, including the VCCC, treat it as native, some as possibly native or probably alien. This species has a coherent European distribution (Jalas & Suominen 1972-1994), reaching the Channel coast wherever there are calcareous soils, though Lambinon et al. (2004) describe it as rare other than in northern Picardy and sometimes occurring as an adventive. In Kent, it occurs also in adjoining semi-natural grassland, albeit in open areas caused by rabbits, but since extensive conservation action has failed to preserve it in such a habitat (T. C. G. Rich, pers comm.), it seems more likely that it has spread from the railway to the grassland, rather than the other way around. With a first date of discovery as late as 1947, its discovery in a railway cutting in Bedford- shire, its presence in a similar habitat in Kent, together with the fact that in Europe it is described as an annual of open ground, all point overwhelmingly to a recent arrival. Decision: alien (neophyte). An example of the distribution being consistent with its European range, with an early date of discovery, but alien status likely SCROPHULARIA SCORODONIA There is no record of cultivation, with the first record in the wild 1689 (Jersey, and 1712 in Cornwall, St Ives). It is a species of open D. A. PEARMAN disturbed ground. Spatially fairly widespread and very dynamic, spreading fast; its persistence in any one site is not known. It has no recorded use, and Watson treated it as native, but it was not covered in Dunn. Recent floras have all treated it as native. The distribution in Britain is a quite consistent extension of its Atlantic range (W France, Spain, Portugal, Madeira, Azores, and N.W. Morocco) In 1950 (Clapham ef al. 1952) it was confined to Cornwall, Devon and Jersey, and naturalised in Glamorgan. By the New Atlas (Preston et al. 2002) it had colonised the S Wales coast to Pembrokeshire, with an outlier in Cardigan, reached Dorset (3 sites) with further outliers further east. It is difficult to escape the conclusion that if one looked at its distribution now, in 2006, knowing nothing of the past, it would look like that of a native, spreading from west to east in a coherent pattern, and recent warm years may be playing a part in this spread. Yet we know it has spread from a few sites, all ports. Davey (1909) gives many sites for Cornwall, though one has the feeling many of these are ruderal; certainly when I moved to Cornwall I was struck by the impression of a fast-colonising weed. The Cornish origin 1s clearly from a number of loci and even today is concentrated around four ports (see the map in French et al. 1999). In Devon it had, before being obscured by later records, an extraordinary clumped distribution around Kingsbridge (Ivemey-Cook 1984). It is now spreading fast (as an alien might), and appears to have no semi-natural niche, that is, though it is a perennial, it does not occur in perennial closed communities. Decision: alien (neophyte). A species in every recent RDB (and protected under the Wildlife and Countryside Act), which is in fact a recent arrival GNAPHALIUM LUTEOALBUM In cultivation 1633, first record in wild 1690 (Jersey, but first British 1802, Cambs and first coastal in Norfolk 1882). It is a species of open disturbed ground. Spatially very rare and scattered, previously declining, but now spreading (Clement 2004); it shows little persistence unless it has continual disturbance. Watson treats it as a casual, but Dunn as a native, with caveats. All recent floras treat it as ; native. STATUS OF DOUBTFULLY NATIVE SPECIES 283 It is found throughout Europe (Hultén & Fries 1986), though the few more detailed maps that I have seen show a very scattered and local distribution (e.g. Haeupler & Schonfelder 1988). In Holland it is colonising new sand-flats on the coast (Mennema ef al. 1980). It is a widely naturalised in the world, especially in warmer climes (Clement 2004), and its true range is difficult to decide. Britten (1899) sets out the early English records, giving the first as from Cambs, reported by the finder as “indubitably wild [that is, by the conventions of the time, native], in the road, far from any house”!! He covers the now vanished Breckland sites, but does not mention the Norfolk coastal populations, which also were reported as “native, remote from _ houses, uninfluenced by agricultural operations and out of the track of the ordinary tourist” (Saunders 1899). There seems little doubt that it is spread by winds and by birds (Clement 2004), and that the seed is persistent, for the current Norfolk sites are in the historic locality, but grow in pits dug in the 1970s and 1980s for natterjacks. There are even more recent records since the New Atlas. The fact that this “serious pest’ (Clement 2004) might be arriving by means other than by man clouds the issue of native or alien status, though there is no doubt it is a recent and repeated arrival, and is categorically unworthy of special protection. Decision: alien (neophyte), with some reservations over the mode of arrival. A cornfield weed, potentially an archaeophyte, but with no evidence yet to support a long sojourn here MELAMPYRUM ARVENSE Never in cultivation, first record in wild 1716 (Norfolk, Norwich) It is a weed of arable habitats. Its distribution is rare and scattered, and it is declining dramatically, though this now masked by conservation actions. It is persistent only where it has spread out of its arable habitat into adjoining bare ground. Watson treats it as a casual or a colonist and Dunn as an alien. Recent floras treat it as native, with some reservations in Stace and Clement & Foster. It is absent from much of western Europe, being frequent in central and _ southern Germany (Haupler & Schonfelder 1988), and widespread east and south of Paris, with outliers in the lower Seine (Institut Floristique Franco-Belge 1978-2001), but rare further west (des Abbayes 1971). Therefore England stands away from the main range. For such a showy plant its first date in the wild is late, especially as it was locally reviled for tainting the wheat. Yet elsewhere in England, in Dorset, it was found as a constituent of smoke-blackened thatch dated to the mid-eighteenth century, suggesting a wider distribution. It is treated as an archaeophyte in the Czech Republic, Poland and Finland, and possibly so in Germany. So there is no question, it seems, as to its status, merely whether it is an archaeophyte or a neophyte, and in the absence of any archaeological evidence and mention in the literature, it seems safer to leave as a neophyte for now. Decision: alien (neophyte, though possible archaeophyte). A late discovery of weedy habitats that has been espoused as a native, possibly to justify retrospectively conservation action FUMARIA REUTERI Never cultivated, first record in wild 1904 (Cornwall, Penryn). It is a species of open disturbed ground. No _ coherent range, declining, with persistence only in allotments, where it is aided by conservation action. Not covered by Watson or Dunn, but treated by all recent floras as native. This species is almost confined to Spain and Portugal, with only one or two records for France, and an historical total of 14 widely scattered sites in England. There is a taxonomic problem here, in that some authorities attribute the English records to the almost endemic subsp. martinii, although Lidén (1986), in Flora Iberica, treats them as synonyms. It seems to be extinct in Cornwall, and as far as is known it is now found only in the Isle of Wight, but at both of these sites it occurs only in allotments. Although it is part of a critical and inconspicuous genus, because of this totally artificial habitat and its lack of contiguous distribution, a late arrival is strongly suggested. This may be contrasted with F. muralis subsp. boraei, one of the group of species that includes F. bastardii, F. capreolata, F. occidentalis and F. purpurea. That species also has a late first date in the wild (1860, presumably reflecting the critical nature of the genus), but, like its congeners, has a niche not 284 D. A. PEARMAN only in arable fields, but also in scrub-edge, hedges and vegetated cliffs. In addition it has a coherent distribution from Spain and Portugal, up the west coast of France to Britain and Ireland. This contrasts with the strongly arable habitat of F. officinalis and others that we treated as archaeophytes. Decision: alien (neophyte). An extinct species, of uncertain status, with a possible very early first record EUPHORBIA VILLOSA The only record of cultivation found is 1758, and the first certain record is 1834 (see below), though both Clarke (1900) and Druce (1932) cite a record from 1576 “in sylva D. Ioannis Coates prope Batthoniam’. It is a species of woods in Europe, but Bath is the only place in Britain that it was ever recorded from. Watson treated it as an alien or denizen, Dunn did not cover it, and all recent floras treated it as alien. The European range is_ easterly and southerly, reaching N.W. France (Clapham et al. 1987), and indeed des Abbayes (1971) gives a few records from the far south of Brittany. Meusel (1965-1992) shows the records reaching the Loire in France, and these French records as isolated from the main centre, with no records in Germany or the Low Countries. The British record therefore is far divorced from any others. I have no idea at all if the species described by Lobel in 1578 was this, though others have so claimed it. Murray (1896) cites a Mr Foster who records that the plant recorded by Lobel was gathered by Thomas Johnson in July 1634. It does not seem to have been recorded or cited by any other author in the next two centuries. However the (re)discovery by Babington, described by him in Smith & Sowerby (1790- 1814) is worth citing. He describes it as “in great plenty and luxuriance in a little frequented lane to the west of Prior Park, and also in a coppice wood, far from any house, to the east. I have not the slightest doubt of it being indigenous in both of the above as there is not the least appearance of it having escaped from any garden”. It was recorded, gradually diminishing, by many others, until the last record in 1941 (Roe 1981). The answer must be academic, other than to those who tabulate the extinctions of native plants, but the evidence for native status seems extraordinarily sketchy. Decision: alien (neophyte). A species treated as native, though we had reservations which now look more sustainable SYMPHYTUM TUBEROSUM In cultivation in 1596, with the first record in wild 1777 (Edinburgh, Water of Leith). It is not included in the NVC but it seems to be a part of woodland vegetation. It is widely distributed, dynamic, increasing fast, and persistent. It is occasionally grown in gardens. Watson treats it as a native, but with caveats re England and part of Scotland; it is not covered in Dunn. Recent floras all treat it as native. The European picture is very interesting. Meusel (1965-1992) shows its range from northern Spain, across southern France to the Black Sea. The German Atlas (Haeupler & Schonfelder 1988) shows it as present only in the farthest south. It is absent from the Low Countries, and present in France only in the centre and south — elsewhere it occurs as an adventive (e.g. des Abbayes 1971). Perring & Walters (1962) treated this probably native only in Scotland, and we have followed this, although the recent Atlas (Preston et al. 2002) contains so many more records, including many more in Scotland too, though some of these may be due to past under-recording. None of the Scottish floras cover this species in any depth, and I have traced only a reference in Watson (1847-1859) citing Gordon (1839) who doubted its native status. Interestingly a later Flora of Moray (Burgess 1935) also calls it alien there. For Cumbria, Halliday (1997) mapped it as native, but mentioned doubts over native status. For Northumberland Swan (1993) also mapped it as native, with a first date of 1820, but his predecessor (Baker & Tate 1867) did not show any records, nor does Swan list any other records between 1820 and 1893. In view of this and its rapid and recent spread in England and Wales, the hallmark of an alien species, together with its continued spread in Scotland, it seems very likely to have been an introduction in the mid-seventeenth century, a conclusion echoed in Braithwaite et al. (2006). Decision: alien (neophyte). A species treated as a native, but about which I now have strong reservations LIMOSELLA AUSTRALIS Never cultivated, with a first record in the wild 1897 (Glamorgan, Kenfig). It seems to have a particular niche in upper salt-marshes. Its STATUS OF DOUBTFULLY NATIVE SPECIES 285 distribution is very limited as a presumed native, but it is stable and persistent. It was not covered in Watson or Dunn, and recent floras have all treated it as native. This species is one of the small group of British species that is not found elsewhere in Europe, though it is widespread elsewhere, particularly in the southern Hemisphere and there is a possibility that its arrival is a recent natural dispersal, or that its presence was overlooked before. It has a very late discovery date, but is very inconspicuous, and appears, in Britain, to be in a finite ecological niche. However Jones (1999) has written persuasively of the possibility of its arrival as a contaminant of ballast, and this seems to me the most likely explanation of its origin, though it does grow with Eleocharis parvula, looking just as native, and that species has only been known in Wales since 1980. Decision: native or alien. A plant previously considered alien where recent work has indicated a niche in semi- natural vegetation VALERIANELLA ERIOCARPA In cultivation 1821, first record in wild 1845 (Worcs, but first in Dorset 1874). Not included in the NVC, but see Pearman & Edwards (2002). Rare, scattered and declining as a casual but stable and confined to chalk and limestone as a presumed native, where it is persistent in open semi-natural vegetation. Use, it is meant to have been one of the species used as “Italian” cornsalad. Watson treated it as a colonist and Dunn as an alien, together with all recent floras. It has no real continuous distribution away from the Mediterranean, or indeed a home even there. On the other hand we suspect nobody has thought about it, and it has always been a critical species! Watson and Dunn, and indeed the twentieth century floras, would probably have thought of it as another weedy species of open habitats, and not known of its cliff-top sites. The work by Pearman & Edwards (2002) seems to show that it is a coherent part of the early-flowering cliff-verge calcareous coastal community (as is Gastridium ventricosum), occurring in communities CG4a, MC5d and MClla (Rodwell 1991-2000), and work subsequent to that paper (Pearman & Edwards 2007) adds sites on the limestone in Devon and Caernarvon, in addition to those already described in Dorset and the Isle of Wight. Decision: native or alien. DISCUSSION THE ROLE OF THE NATURE CONSERVATION MOVEMENT The twentieth century growth of the nature conservation movement has _ brought developments that would have surprised our earlier botanists, notably the concentration of resources on species perceived to be native. Again this is dealt with in more depth in Preston et al. (2004), but the result has been that only species regarded as native or archaeophyte have been included in the current Red Data List (Cheffings & Farrell 2005), the recent Red Data Book (Wigginton 1999), or designated as nationally scarce (Stewart et al. 1994). The BAP list similarly is comprised exclusively of perceived natives, and though the country conservation organisations have now treated archaeophytes as “honorary natives” for conservation purposes, and similarly those species designated as “native or alien” in the new Atlas, this treatment has not been extended to those we designated, often for the first time in a century or more, as neophytes. Of course resources themselves are scarce, but this, if I may describe it, obsession with native status, which extends in the opposite direction to blaming aliens for many of the ills facing our native flora, is surely misplaced and simplistic. I would like to extend the parameters of species worthy of protection to those species which are an integral part of the mosaic which is being valued and conserved — thus, amongst others, one might preserve Fritillaria meleagris, Lonicera xylosteum, Salvia pratensis and perhaps Teucrium chamaedrys as being part of a valued semi- natural habitat; Althaea hirsuta, Anisantha madritensis, Galium parisiense, Holosteum umbellatum, Teucrium botrys and_ possibly Petrorhagia prolifera as species very persistent in open, species-rich, therophyte communities; Melampyrum arvense and Rhinanthus angustifolius as part of the cornfield weed mosaic we now value (and possibly potential archaeophytes too). I appreciate this is more subjective than the natives “good”, aliens “bad” approach, but I do feel that species that have that cultural resonance, act in harmony with other species that we value in their habitats and have a long association in those sites, could well be included in conservation efforts. This would be all the more possible since almost all conservation efforts are perforce concentrated on the management of the habitat rather than the individual species therein. Furthermore the list would not be long. 286 D. A. PEARMAN Conversely there must be some effort by the conservation movement to absorb new evidence, to review S.S.S.I.s notified for their botanical interest, and amend them if necessary. I see no need, for instance, for S.S.S..s for Equisetum ramosissimum or Cerastium brachypetalum on old railway sites, unless they fit the suggested guidelines above. A PLEA FOR A MORE HOLISTIC APPROACH There has, then, been relatively little discussion in the round on the possible status of plants in Britain in the century or more since Watson and Dunn; in fact, other than in Lousley (1953) and Webb (1985), I have been unable to find any substantial work in that period, and neither appears to have had any influence on later writers, though the latter is often cited! National floras since 1950 have tended to take a much more simple approach than Watson, for reasons I cannot pin down, though Preston et al. (2004) detect the influence of Godwin’s work on the history of the British flora (Godwin 1956); this influence would of course extend to the treatment of archaeophytes rather to neophytes. A further interesting point might be that in Clapham et al. (1952 etc), there was, for the first time in well over a century, one acknowledged national flora, which was followed by all of their generation. Local Floras have often continued to look at the position in their area, rather than stand back and look at the national or even European picture (Halliday (1997) is a_ notable exception). In these local floras and other papers the role of the optimist was able to flourish. In addition to White (1912) mentioned above, the illustrious names of Marshall (Hanbury & Marshall 1899), and the late Francis Rose head a list of consistent pleaders for native status for favoured species, along with a host of Flora writers. An extreme illustration of optimism comes from Cambridgeshire (Anon 1960), where the finding of a patch of Aster salignus [actually A.novi-belgii|, previously under suspicion as a garden escape at Wicken Fen, at Fowlmere “many yards from the nearest house means that its status must be carefully reconsidered”. As in the concept of Archaeophytes it does not seem, in the main, that British writers have used the slowly-growing corpus of grid-based map floras from Europe and their supporting literature, and have not obviously been aware of both the developments in the world of archaeobotanical research and of interpretation of the names used in medieval gardening (or even that of the 16th, 17th and early 18th centuries), and even of the existence of European works on alien floras that are becoming available. What case studies there have been have tended to have been restricted to individual species, which may then have been examined not only in isolation to others, but all to often to a limited range of criteria. Inevitable though this might be, it does not seem a fruitful way forward, and that is why, for the New Atlas, and therefore as the basis of this paper, we chose to examine a large number of species and to try and apply the same criteria to all. Of course I appreciate that for many species there can never be certainty, though for neophytes the imponderables must usually be less than for archaeophytes. However an awareness of the sources outlined in this paper must still have more to contribute — witness the case of Fritillaria, fairly clear already (to those who were prepared to look), that has become clearer through examination of Scandinavian work and our own county floras. There must be more work to do in unravelling the history both of species treated here (and in the available spreadsheet) as neophytes and of others not yet fully considered. In alphabetical order I would like to investigate, for instance, Allium scorodoprasum, Crepis foetida, Ericas in Ireland, Lepidium latifolium (at least away from the east coast), Scrophularia umbrosa, Sorbus domestica and Stachys germanica. Other species covered in our initial list might well turn out to be archaeophytes if more historical evidence becomes available. Webb (1985) has the magisterial status that I am sure I lack. I was reading British Wildlife last month, and came across a comment that Sisyrinchium bermudiana might well be native in the Wye valley, “remote from any houses” (Peterken & Tyler 2006). It reminded me of Webb’s closing comments “The various arguments must be weighed against each another as fairly as possible, and the conclusion adopted which, even if rather improbable, seems less improbable than the alternatives. Who is best qualified to do this weighing? ....Detailed local knowledge is _ often invaluable, but all too often its value is eroded by local patriotism. There is a _ curious emotional bias, which I have found very widespread (and from which I may not entirely be free myself) which favours native status for STATUS OF DOUBTFULLY NATIVE SPECIES 287 an attractive plant or for the botanist’s home county. For this reason a cool assessment by an outsider may be more reliable.” I hope this paper sets the scene of that re-appraisal; what is really needed is a series of case studies on all these contentious species, to enable the criteria for each to be aired and placed in context. ACKNOWLEDGMENTS The principal influence on the information presented here, and in the database wherein lies all the underlying research has been Chris Preston. I always imagined we would write this paper together, as a complimentary companion to that on archaeophytes, but his workload has not allowed this. I am very grateful for all his research which has been made available to me and for his comments, without which this paper could not have been written, though I must take responsibility for all the interpretations. | am particularly grateful too to Kevin Walker for comments and help with accessioning papers, to Andy Byfield and Alex Lockton, and to French colleagues for making their works available to me. 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Vegetation ecology and Population biology of Fritillaria meleagris L. at the Kungsangen Nature Reserve, Eastern Sweden. Acta Phytogeographica Suecica 73. Uppsala. (Accepted November 2006) Watsonia 26: 291—302 (2007) The last thirty five years: recent changes in the flora of the British Isles K. J. WALKER* CEH Monks Wood, Abbots Ripton, Huntingdon, Cambs. PE28 2LS ABSTRACT In 1970 Max Walters predicted a number of changes that have taken place in the British and Irish flora over the last 35 years. Although few species became nationally extinct during this period, losses at the local scale were dramatic, particularly in S.E. England where land use pressures were greatest. Despite these losses the British flora has become increasingly diverse due to the introduction, natural- isation and spread of alien species. Research into their habitats, modes of dispersal, and competitive inter-actions with native species has _ provided valuable ecological insights. Hybridisation events have produced some of the best studied examples of hybrid speciation. In the future, such naturalisation events may also provide important evidence for major environmental perturbations such as climate change. KEYWORDS: aliens, diversity, extinction, hybrid- isation, naturalisation, neophytes, urban habitats. INTRODUCTION In 1970 Max Walters concluded The flora of a changing Britain conference by predicting a number of changes that were likely to take place in the British flora over the subsequent twenty-five years (Walters 1970). Although brief, his paper dealt with floristic richness, aliens, extinctions, plant conservation and land abandonment. The aim of this paper is to review three of these predictions in view of the major changes which have taken place: (1) floristic richness has increased as a result of man’s activities; (2) the number of national and regional extinctions has continued to increase; and (3) much has been learnt from the naturalisation of alien species. In addressing these three predictions I have intentionally drawn heavily (but not exclusively) on research published within this journal, especially the review by Brenan (1982) which deals with many of the same topics. Wherever possible, I have also referred to changes in Max’s home *e-mail: kwal@ceh.ac.uk county of Cambridgeshire (v.c. 29). I do not deal with changes in plant conservation, land use, or the ten activities he proposed for the society as these now form the basis of much of the society’s current work (e.g. monitoring common species, research on rare species, publication of handbooks). 1. FLORISTIC RICHNESS WILL CONTINUE TO INCREASE AS A RESULT OF MAN’S ACTIVITIES As Max predicted species invasions have exceeded extinctions by an order of magnitude leading to a dramatic net increase in the number of plant species recorded in the British Isles (Fig. 1). Despite localised losses only 21 native species have gone extinct nationally since records began in the sixteenth century (Stace 2002). Nine of these species have been lost in the last 35 years, although some are either dubiously native (e.g. Galeopsis sege- tum) or are likely to have been over-looked (e.g. Epipogium aphyllum) (Table 1). During the same period an equal number of species have been discovered or newly described of which only two are of questionable origin (Cystopteris diaphana, Serapias parviflora) and one (Epipactis youngiana) may be a recently evolved ecotype (Hollingsworth er al. 2006). In comparison to these rather modest changes to the native flora, there has been a dramatic increase in the number of naturalised aliens and casuals (Fig. 1): by the middle of the 1950s the number of recorded casuals exceeded the number of natives whereas there were around 600 naturalised aliens (30% of the native flora). By 2000 the number of casuals and naturalised aliens had increased to make up 45% and 18% of the entire British flora respectively. . Historically alien plants have arrived unintentionally as escapes from botanic gar- dens or as contaminants in the ballast of ships, grain, animal fodder and waste products of 292 K. J. WALKER Native et eeeee Naturalised aliens 3000 | —— ——Casuals a 2) a) (S) ® (ok 2) — e) i= ® & = =] = 500 0 + T T T 1900 1920 1940 1960 1980 2000 Year FIGURE |. The number of native taxa, naturalised aliens and casuals in the flora of Great Britain according to various sources (Druce 1908; Druce 1928; Dandy 1958; Kent 1992; Kent & Stace 2000). “Native’ includes agamospecies such as Hieracium, Rubus and Taraxacum. Adapted from Stace (2002). TABLE 1. NATIVE SPECIES DISCOVERED OR KNOWN TO HAVE BECOME EXTINCT IN BRITAIN (INCLUDING ISLE OF MAN) SINCE 1970. UPDATED FROM STACE (2002). SPECIES IN THE CRITICAL GENERA ALCHEMILLA, EUPHRASIA, HIERACIUM, LIMONIUM, RUBUS, SORBUS AND TARAXACUM ARE OMITTED Species Prrsulast V.C. Comment record (a) Discovered Epipactis youngiana 1976 67 Probably only an ecotype of E. helleborine Senecio eboracensis O79 63 Derived from spontaneous hybrid Crepis praemorsa 1988 69 Utricularia stygia 1988 Oy) Serapias parviflora 1989 2 Native status uncertain Callitriche palustris 2000 99 Cystopteris diaphana 2000 2 Native status uncertain Carex salina 2004 105 (b) Extinct Arnoseris minima 1971 24 ~~ Probable archaeophyte. Re-introduced Bromus interruptus 1972 29 Probable neophyte. Re-introduced Schoenoplectus 1972 59 Only native on Jersey. Re-introduced pungens Galeopsis segetum NOs) 49 Probable archaeophyte Saxifraga rosacea 1978 49 Still occurs in Ireland subsp. rosacea Crepis foetida 1980 8) Probable archaeophyte. Re-introduced Epipogium aphyllum 1986 22 Unconfirmed records since 1990 Neotinea maculata 1986 a Occurs in Ireland. Formerly Isle of Man Senecio eboracensis 2000 - 61 Possibly extinct? Luzula pallidula is included as extinct by Stace (2002) but there are a number of confirmed records from Holme Fen in Huntingdonshire (v.c. 31). . RECENT CHANGES IN THE FLORA OF THE BRITISH ISLES 293 TABLE 2. CHANGE IN THE NUMBER OF TAXA RECORDED IN BERKSHIRE (V.C. 22). ADAPTED FROM CRAWLEY (2005) Druce (1897) (1968) Native & archaeophyte 820 883 Neophyte 300 629 Apomictic 63 84 Subspecies & varieties 400 38 Hybrids 70 99 clothing manufacture (skins, wool, tan-bark) (Lousley 1953). Although introductions via these agencies declined during the second half of the twentieth century (Shimwell 2006), the numbers originating from horticulture, either as escapes or outcasts from gardens, have in- creased dramatically. For example, the New Atlas showed that eight of the 10 most rapidly Spreading species since the 1960s are neophytes which were originally introduced as garden plants (Buddleja davidii, Cotoneaster simonsil, Laburnum anagyroides, Linaria purpurea, Lysimachia punctata, Prunus cerasus, P. laurocerasus, Syringa vulgaris) (Preston et al. 2002). Changes at the vice-county level show a similar pattern. Berkshire (v.c. 22) has one of the best studied floras in the country, parti- cularly with regard to alien species, and this 500 a (a) : £ 400 - ° o. & .o) Se. e) Os ea a2 e 6 % Yo ° Oo © PaeDOy < = (crn) - eo) OY 6 fo) © 200 - roe) cB © © 0 00M O > PEP B 0 O = 0° @ 9° ® nose | ye za Se re) © fe) ie) co O (e) (@) i —4) 0.1 1.0 10.0 100.0 0 . =1 Population density (ha _) Bowen Crawley % change (2005) 1897-1968 1968-2005 955 +8 +8 710 +110 12 184 33 +119 Al -9] +8 181 +4] +83 has allowed an assessment of changes in floristic richness over the past 100 years (Table 2; Crawley 2005). Whereas the number of native and archaeophyte additions has been rather modest there has been a dramatic increase in the number of recorded neophytes (Bowen 1968). Similarly, critical and hybrid taxa have been much better studied since 1968 whereas the vast majority of infraspecific taxa formerly recognised by Druce (1897) are not included in the modern floras of Bowen and Crawley. Based on his experiences in Cambridgeshire Max Walters predicted that urban areas would accrue more alien species than rural areas and therefore, that urbanised 10-km squares would be the richest. However, unlike continental Europe, British urbanised tetrads are not significantly richer overall than their rural 150 ; (b) — O e) sy oO f® 100 - § ae © Q O 2 | saa oO O J Dp © OG, o ® ony za O BGS Sd “6 | a OG BB Og. 5 £28 és or S60 | 0.1 1.0 10.0 100.0 Population density (ha’') FIGURE 2. The relationship between human population density (displayed on a log scale) and (a) the number of natives and archaeophytes and (b) neophytes and casuals recorded in Cambridgeshire parishes (v.c. 29) between 1987-2000 (n=151). Human population densities were calculated from Population Census Returns for 2001. 294 K. J. WALKER counterparts, although they do support a greater number (and proportion) of introduced alien species (Roy et al. 1999). The results for agriculturally ravaged Cambridgeshire were as predicted by Max with more densely populated (and therefore urbanised) parishes tending to be marginally more species-rich (R°=4:2%, F=6:5, p <0:-05) whereas differences in the number of natives and archaeophytes were not significant due to large variation in numbers of species between parishes (R°=1:9%, F=2-9, p=0-09; Fig. 2a). In contrast, urbanised parishes have significantly more neophytes and casuals with several of the ten richest parishes being almost entirely urban (R°=16:9%, F=30°3, pp <0: 001 Fig. 2b). The slight discrepancy between Cambridgeshire and the national picture (1.e. more species in urban areas) may be due to local factors. Some urban parishes in the county have been extraordinarily well recorded by resident botanists (e.g. Cambridge, Chester- ton, Ely, Whittlesey) whereas the majority of rural parishes are relatively species-poor due to the intensity of arable farming in the county. 2. NATIONAL AND REGIONAL EXTINCTIONS WILL CONTINUE AT AN ALARMING RATE As discussed above relatively few species have gone extinct at the national level and, with the possible exceptions of Bromus interruptus, which may be a recently evolved neo-endemic (Rich & Lockton 2002), and Senecio eboracensis, none have become — globally extinct. Of the nine species that have disappeared since 1970 (Table 1) five were probably originally introduced (Arnoseris minima, Bromus interruptus, Crepis foetida, Galeopsis segetum, Schoenoplectus pungens; but see Rich & Lockton (2002) and Rich & Pryor (2003) for alternative views), Saxifraga rosacea and Neotinea maculata still occur in Ireland and there are unconfirmed records for Epipogium aphyllum in the last decade. In contrast, Senecio eboracensis is probably the shortest-lived addition to the British flora, having possibly gone extinct within 17 years of its discovery in 1979 (Abbott et al. 2005). In contrast to these rather modest changes at the national level, there has been a dramatic loss of species at the regional and county level since the nineteenth century. Twenty-three native species have gone extinct in the most intensively populated region of lowland England which includes all vice-counties to the east of a line running from Poole in Dorset to Goole in S.W. Yorkshire (‘the Poole to Goole line’; Table 3). The vast majority are Northern (Boreal) species at the southern edge of their range in lowland England that disappeared following the widespread loss of lowland heaths and bogs (e.g. Diphasiastrum spp., Huperzia selago, Gymnocarpium dryopteris, Listera cordata, Pseudorchis albida, Pyrola media, Scheuchzeria palustris, Selaginella selaginoides, Trientalis europaeus, Vaccinium vitis-idaea). Others have been lost due to drainage of fenland (Tephroseris palustris), hybridisation (Schoenoplectus _ triqueter), ploughing of calcareous grassland (Cirsium tuberosum) and changes to aquatic habitats (Elatine hydropiper, Luronium natans) or simply because their extreme rarity in S.E. England has made them highly susceptible to other factors (Asplenium obovatum, A. viride, Genista pilosa, Mertensia maritima, Orobanche alba, Vicia orobus). Species which still occur in S.E. England but have suffered dramatic declines are also listed in Table 3. These show a similar pattern to extinctions with the majority of species associated with rivers and wetlands (Damasonium alisma, Leersia oryzoides, Potamogeton praelongus, Teucrium scordium), infertile grasslands and heaths (Antennaria dioica, Gentianella campestris, Orchis militaris, Potentilla neu- manniana) and lowland bogs and mires (Carex limosa, Liparis loeselii, Viola persicifolia, Utricularia intermedia). The loss of species at the county level has been much greater than at the national or regional scale (Walker 2003). Fig. 3 shows the average decadal rate of extinction (actually the number of last records) using the most recent information for eighteen vice-counties in England (data available from the author on request). This shows a gradual increase in the number of extinctions from less than one species per decade at the start of the nineteenth century to 3 species per decade by 1850 and 4 species per decade by 1900. Numbers were more variable during the first half of the twentieth century, but were probably around 5 species per decade up to 1950 increasing to 6 species per decade in the 1960s and 1970s. The apparent decline in the 1980s is undoubtedly an underestimate as more time is usually needed to declare a species as extinct. Although the figures for the nineteenth century are less reliable due to lower recording intensities, they RECENT CHANGES IN THE FLORA OF THE BRITISH ISLES 295 TABLE 3. EXTINCT AND DECLINING SPECIES IN SOUTH EAST ENGLAND! (a) Extinct in S.E. England Loss (b) Marked decline in S.E. England? Extant Loss Tephroseris palustris 26 Gastridium ventricosum 5 91 Huperzia selago 24 Potamogeton praelongus 5 2 )3) Gymnocarpium dryopteris 13 Damasonium alisma 3 46 Genista pilosa 1 Gentianella campestris 3) 46 Pseudorchis albida 7 Antennaria dioica 1 34 Schoenoplectus triqueter 6 Lactuca saligna 3 33 Elatine hydropiper >) Cynoglossum germanicum 3) 30 Listera cordata + Teucrium scordium l 20 Epipogium aphyllum 3} Atriplex pedunculata 17/ Selaginella selaginoides 3 Utricularia intermedia s.1. 177 Asplenium viride Dp Liparis loeselii D 16 Luronium natans 2 Orchis militaris 3 16 Orobanche alba 2 Alchemilla glabra 4 14 Scheuchzeria palustris 2 Leersia oryzoides 3 14 Asplenium obovatum 1 Viola persicifolia 3 14 Cirsium tuberosum 1 Campanula patula 3 3) Diphasiastrum alpinum 1 Vicia bithynica 5 13 Diphasiastrum complanatum 1 Carex limosa 3 12 Mertensia maritima 1 Erodium maritimum 3 10 Pyrola media 1 Potentilla neumanniana 5) 10 Trientalis europaeus 1 Lotus angustissimus 5 9 Vaccinium vitis-idaea 1 Salicornia nitens 3 9 Vicia orobus 1 Hypochaeris maculata 5 8 Includes vice-counties 10-32 & 53-56. *Species recorded in <5 10-km squares between 1987 and 2000 (Preston et al. 2002). ‘Loss’ is the number of 10-km squares in which a species was recorded prior to 1987. A further 65 species occur in <5 10-km squares in S.E. England but have been excluded because they are either (a) very rare (n=21), (b) rare but showing only localised declines (n=23) or (c) northern/western species at the extreme edge of their range in S.E. England (n=21): (a) Apium repens, Artemisia campestris, Calamagrostis stricta, Cephalanthera rubra, Cyperus fuscus, Eriophorum gracile, Gnaphalium luteoalbum, Goodyera repens, Ludwigia palustris, Orchis simia, Orobanche artemisiae-campestris, Petrorhagia nanteuilii, Phyteuma spicatum, Polygala amarella, Potamogeton nodosus, Salicornia obscura, Scleranthus perennis, Scilla autumnalis, Senecio paludosus, Stachys germanica, Thymus serpyllum, Veronica spicata, Veronica verna. (b) Alisma gramineum, Atriplex longipes, Bupleurum baldense, Carex depauperata, Carex filiformis, Clinopodium menthifolium, Eleocharis parvula, Gentianella ciliata, Lobelia urens, Luzula_pallidula, Maianthemum bifolium, Najas marina, Ophioglossum azoricum, Ophrys fuciflora, Orobanche caryophyllea, Pilosella peleteriana, Polygonum maritimum, Pulmonaria obscura, Selinum carvifolia, Seseli libanotis, Thlaspi perfoliatum. (c) Allium scorodoprasum, Asplenium marinum, Blysmus rufus, Carum verticillatum, Dactylorhiza purpurella, Equisetum variegatum, Euphorbia portlandica, Festuca altissima, Fumaria_ purpurea, Gymnocarpium robertianum, Hymenophyllum tunbrigense, Juncus acutus, Juncus filiformis, Lavatera arborea, Melica nutans, Phegopteris connectilis, Rosa mollis, Rubus saxatilis, Salix myrsinifolia, Sibthorpia europaea, Sparganium angustifolium, Stellaria nemorum. suggest an overall increase in the rate of extinction during the twentieth century, especially since the 1960s. As can be seen from Fig. 3, rates in Cambridgeshire were generally higher than average throughout this period, particularly during the first half of the nineteenth century when many infertile grassland and bog species (e.g. Drosera intermedia, D. anglica, Herminium monorchis, Narthecium ossifragum, Ophrys_ sphegodes) were lost as a result of parliamentary enclosure © (Preston 2000). Rates approached one species a year (10 species per decade) for the majority of the twentieth century, particularly after the 1950s, when species of infertile semi-natural habitats continued to decline at an alarming 296 K. J. WALKER 12 . @ Allcounties aR RN arc Cambr idgeshire \ Comal oe D> 10 - tt i | Sc ae a) - i o | i ®o® 8+ | TS te | ( Known from a single site in Leicestershire up till 1934. * There is an earlier record for 1838. * The first record was from Guernsey. The first record in England was from S Essex in 1974. widely naturalised species are either highly competitive woody or thicket-forming species capable of excluding native vegetation (e.g. Fallopia japonica) or rapidly maturing species that succumb to competition during secondary succession but spread rapidly via disturbed niches (e.g. Epilobium ciliatum, Senecio squalidus). Some of these invaders display higher growth rates than their native or introduced congeners. For example, stems of Elodea nuttallii elongate faster than those of E. canadensis, and this has been shown to cause displacement of the latter, formerly invasive Species, in some sites (Simpson 1984, 1990). Similarly, the rapid development of leaves in Lemna minuta can exclude other native floating aquatics (Leslie & Walters 1983). In ruderal habitats, the dramatic spread of Epilobium ciliatum may be due, in part, to a faster growth rate and longer flower period than native willowherbs (Preston 1988). The lack of natural pests and pathogens within the introduced range may also provide exotic species with a competitive advantage thereby making them more invasive (the ‘enemy release hypothesis’; Keane & Crawley 2002). Perhaps the most notoriously invasive plant in the British flora, Fallopia japonica var. Japonica, provides a good example. Plants in Japan suffer damage from a range of invertebrate pests which attack rhizomes, stems and leaves. No such pests have been found on British plants and no doubt this has aided its dramatic spread following its escape from cultivation in 1886 (Bailey & Connolly 2000). HABITAT The habitats occupied by naturalised species can provide fascinating ecological insights as well as providing novel conditions in which to study interactions with native congeners (e.g. competition, hydridisation). The vast majority of alien plants occur in man-made habitats (e.g. waste ground, railway lines, walls, urban sites) close to habitation (‘the 100 m rule’; Crawley et al. 1996). These habitats provide disturbed, and often drought-stressed niches (e.g. cinder tips, clinker of railway lines, asphalt, quarries, rubble, wasteland) similar to conditions occupied in the native range. Some of the best examples are primary colonists of volcanic rocks (e.g. Senecio squalidus, Fallopia Japonicus var. japonicus) and river shingles and gravels (e.g. Epilobium brunnescens). Buddleja davidii is a perennial shrub native to central and western China where it typically occurs as dense thickets on shingle beaches adjacent to rivers (Owen & Whiteway 1980). Its spread in southern Britain was one of the most dramatic of the twentieth century and 298 followed the sudden availability of lime-rich rubble created by bombing during the 1939- 1945 war. Its ability to thrive on compacted, drought-prone soils combined with prolific seed production (>1 million seeds per plant) has allowed it to colonise a wide variety of ruderal niches (e.g. railway sidings, quarries, stone structures) and as a result it is now one of the most familiar weeds of urban and waste areas (Clay & Drinkall 2001). Similarly, Epilobium brunnescens has spread throughout much of northern and western Britain where it is now common on moist, gravely substrates similar to its native riverbed habitat in New Zealand (Kitchener & McKean 1998). Urban niches also provide suitable conditions for more southerly species such as the subtropical plant Conyza sumatrensis which is confined to arid, sun-baked niches such as pavement cracks (Wurzell 1988). In contrast, it’s more temperate cousin Conyza canadensis has spread further and more rapidly along railway- lines since the 1939-1945 war (Salisbury 1961). INTRODUCTION AND SPREAD Although some _ naturalised aliens were introduced unintentionally the vast majority, including many pernicious plant invaders, were brought in as garden ornamentals (Crawley et al. 1996). Their subsequent spread has long fascinated botanists and as a result many have been studied in great detail (e.g. Elodea spp.., Fallopia spp., Senecio spp.). The time lag between escape into the wild and invasion can be surprisingly long given how widespread some species eventually become. For example, Epilobium ciliatum was known for over 40 years from a single site in Leicestershire before it began its dramatic spread throughout southern England in the 1930s (Preston 1988). However, most plant invasions have been shown to follow a rather generalised pattern with a pioneer phase of scattered primary occurrences followed by a rapid expansion from primary foci (‘infection’ sensu Salisbury 1961) and a final era of consolidation. The naturalisation of Fallopia japonica var. japonica in the British Isles provides the classic example. Its spread began in the early 1900s with initial foci in industrial areas in South Wales but also private gardens, estates, nurseries and botanic gardens from which dispersal by escape, exchange, gift, sale or disposal took place (Bailey & Connolly 2000). K. J. WALKER Indeed today clusters of hybrids with F. sachalinensis (F. xbohemica) often indicate proximity to gardens where it was originally planted (Bailey & Conolly 2000). It spread modestly up till the 1939-1945 war but rapidly thereafter, especially around urban areas, before consolidating its range in the 1970s (Conolly 1977). In contrast, wind dispersed aliens have shown a much more continuous pattern of spread, particularly where colonisation took place along transport corridors (e.g. railway sidings). For example, Epilobium ciliatum showed a more or less even spread after the 1930s reflecting a more natural expansion of an established population rather than introduction by man at different sites (Preston 1988). Although most aliens consolidate their distributions, a small number have sub- — sequently declined. One of the best examples 1s Elodea canadensis: between 1850 and 1880 this North American species spread vigorously in the east of England, where it blocked drains, sluices and watercourses much to the chagrin of college rowers and swimmers in Cambridge (Simpson 1984). However, it quite inexplicably declined in abundance and invasiveness, more latterly due to displacement by E. nuttallii which was first recorded near Oxford in 1966 (Simpson 1990). Spartina anglica, the allopolyploid derivative of S. maritima and the North American S. alterniflora, has suffered a similar ‘die-back’ in some areas due to successional changes (Ainouche ef al. 2004). DISPERSAL MODE Research on the dispersal modes of aliens has identified two distinct groups of successful invaders: tall, competitors that spread largely by vegetative fragments in garden waste and small, short-lived species with high reproductive outputs which are dispersed in soil or as airborne propagules (Hodkinson & Thompson 1997). The most extreme example of the former group is Fallopia japonica var. Japonica. British populations are male-sterile and so spread is entirely by vegetative fragments from a single extraordinarily vig- orous clone (Conolly 1977; Hollingsworth & Bailey 2000). Plants can establish from tiny root fragments dispersed in topsoil or garden waste, making it almost impossible to eradicate or control. The spread of the diminutive rockery plant Veronica filiformis has received less attention but is no less dramatic. Despite RECENT CHANGES IN THE FLORA OF THE BRITISH ISLES 299 setting virtually no seed, it has colonised almost every lowland hectad in less than 70 years, presumably by vegetative fragments transported on mowing machinery. Likewise, the astonishing spread of Spartina anglica along the Western seaboard of Europe has occurred as a result of dispersal of rhizome fragments by sea currents. The spread of Epilobium ciliatum and Senecio squalidus are relatively easier to understand as both produce copious amounts of plumed seed capable of dispersal over large distances. For example, Druce (1927, p.241) famously observed the dispersal of seeds S. squalidus during a railway journey between Oxford and Reading (c. 40 miles): “...the vortex of air following the express train carries the fruits in its wake. I have seen them enter a railway-carriage window near Oxford and remain suspended in the air in the compartment until they found an exit at Tilehurst’”. No doubt many other ‘railway yard’ aliens have been spread in a similar fashion (e.g. Buddleja davidii, Conyza canadensis, Senecio viscosus). Invasion of riparian habitats via waterborne dispersal of seeds, fruits and vegetative propagules have been no less dramatic. Since the 1940s Impatiens glandulifera (incidentally Britain’s tallest annual plant) has spread rapidly along riverbanks throughout the British Isles and now occurs in dense mono-specific stands where it can suppress native perennials such as Urtica dioica (Tickner et al. 2001). Less invasive examples include the dispersal of seeds of Lathraea clandestina, bulbils of Cardamine bulbifera and vegetative fragments of Arenaria balearica along watercourses (Atkinson 1996; Corner 1996; Wallace 2005). The transport of seeds in soils or waste manures (e.g. wool shoddy) has also led to the spread of many species, although relatively few aliens have become widely naturalised via this agency. Possibly the most successful example is Matricaria discoidea which was originally introduced, into Ireland at least, with American corn fed to poultry (Reynolds 1996) but has since spread rapidly in soil attached to animals, shoes and motor vehicles (Hodkinson & Thompson 1997). A more recent invasion has been the spread of Galinsoga quadriradiata in urban areas, initially due to the effects of bombing during the 1939-1945 war, but more latterly in soil amongst horticultural produce (Lacey 1957). HYBRIDISATION AND SPECIATION Occasionally naturalised species hybridise with native congeners or other introduced species. Of the estimated 800 or so angiosperm hybrids known to occur in the UK 87 combinations involve at least one non-native taxon and 26 are hybrids between two or more alien species (Preston et al. 2002). Many originated allopatrically and are therefore unique to the British Isles most notably hybrids between Fallopia and Epilobium taxa. As _ British populations of Fallopia japonica var. japonica are entirely male sterile all seed produced is inevitably of hybrid origin. The most frequent pollen donor appears to be the common garden plant, F. baldschuanica although instances of the hybrid F. xconollyana establishing in the wild have been astonishingly rare given the amount of seed produced annually (Bailey 2001; but see Bailey & Spencer 2003). In contrast, the hybrid with F. sachalinensis (=F. xbohemica) displays many of the invasive tendencies of F. japonica var. japonica and is currently colonising roadsides, waste ground, riversides and railway banks throughout the British Isles (Bailey et al. 1996). Harsh, man-made environments often create novel abiotic conditions in which evolutionary changes can take place (e.g. heavy metal tolerance on colliery waste tips). They can also stimulate allopatric speciation events and new hybrid combinations. Good examples are hybrids between the naturalised willowherb Epilobium brunnescens and _ native E. lanceolatum, E. montanum, E. obscurum, E. palustre and E. parviflorum. All have arisen relatively recently on damp, acid substrates associated with former mining activities (e.g. quarry slopes, granite aggregates, mine wastes, china clay gravels) in Cornwall (Kitchener & McKean 1998; Kitchener 2003a). Native plants are very slow to colonise these habitats and as a consequence primary colonists, such as willowherbs, have ample to time to establish and hybridise (Kitchener 2003b). In addition, the relatively sunny and warm Cornish climate may encourage hybridisation by allowing flowers to remain open longer thereby increasing chances of cross-pollination by insects (Kitchener 2003b). The evolutionary implications of such novel hybridisation events are well documented and provide some of the clearest examples of 300 K. J. WALKER speciation by allopolyploidy. Possibly the best studied is the neoallopolyploid Spartina anglica which originated from hybridisation between the native species S. maritima and East American S. alterniflora following its introduction to Southampton Water in the 1820s. The sterile first generation hybrid, S. xtownsendii, was first observed in 1870 and within twenty years gave rise to the fertile allopolyploid S. anglica following genome duplication (Ainouche et al. 2004). More recent examples have involved Senecio squa- lidus, a diploid ragwort introduced from Mount Etna, Sicily, at the beginning of the eighteenth century (Harris 2002). This species may have originated in Oxford as a stabilised diploid hybrid between two Sicilian species, S. aethnensis and S. chrysanthemifolius (Abbott et al. 2000, 2002; although Sell & Murrell (2006) treat both as subspecies of S. squalidus). Hybridisation between S. squalidus and S. vulgaris 1s relatively rare in the wild yielding highly sterile triploid (=S. xbaxteri) or partially fertile tetraploid hybrids. Segregation from either of these hybrids, or following backcrossing to S. vulgaris, has been shown to produce the ‘York radiate groundsel’ S. eboracensis (Lowe & Abbott 2003) in England and the allohexaploid ‘Welsh groundsel’ Senecio cambrensis in Wales and Scotland (Abbott et al. 1983; Ashton & Abbott 1992). Although reproductively isolated from their parents due to self-compatibility mechanisms and flowering times (Lowe & Abbott 2004) both are currently threatened: the Edinburgh lineage of S. cambrensis was _ recently destroyed by redevelopment (Abbott & Forbes 2002) and York populations of S. eboracensis have declined as a result of over-zealous street cleansing and may be extinct (Abbott ef al. 2005). Introgression of S. squalidus into S. vulgaris subsp. vulgaris has also been implicated in the origin of radiate forms of groundsel, S. vulgaris var. hibernicus, which 1s now a common variety of town and wasteland floras in many parts of the British Isles (Abbott et al. 1992). Senecio squalidus also hybridises with the introduced S. viscosus to form the sterile hybrid S. xsubnebrodensis and it has been suggested that subsequent introgression may have led to the spread of S. viscosus during the twentieth century (Crisp & Jones 1978). CONCLUSION The rather depauperate nature of the British flora has meant that, with the possibility of Bromus interruptus, Senecio eboracensis and a few critical taxa, no species have become globally extinct, and relatively few have been lost in the UK as a whole. However, as Max Walters (1970) predicted this belies real losses at the local scale over the last 35 years, particularly in the intensively managed lowlands of southern England, where land use pressures have been greatest. Here the extinction of species of infertile habitats shows now sign of abating, and it seems likely that many of the species listed in Table 3 could well go extinct within the next 35 years. However, these localised losses have been more than compensated by the discovery of new species, - mainly within critical genera, and, even more so by the introduction and spread of casuals and neophytes. Although Max’s prediction that urban areas will be the most diverse is not strictly true, they are certainly the centres of alien plant diversity in the UK and, as the results from Cambridgeshire show, they may well become the richest areas if we continue to lose native species from the countryside at the same alarming rate. In recent decades many non-native species have been treated as unwelcome additions which will threaten our native flora.. However, British botanists have learnt a great deal from their means of arrival, dispersal, habitats and competitive interactions with the native flora. In addition, the naturalisation of species from all over the globe has provided evolutionary biologists with unique opportunities to study the intricacies of diploid hybrid speciation. Future naturalisation events may also provide evidence of major environmental changes, such as_ climate change, as conditions become more suitable for Mediterranean and subtropical species. ACKNOWLEDGMENTS I would like to thank David Pearman for comments on an earlier version of this paper, Colin French and Ian Bennallick for providing information on extinctions in Cornwall, and Nick Millar for permission to use data on Cambridgeshire parishes. RECENT CHANGES IN THE FLORA OF THE BRITISH ISLES 301 REFERENCES ABBOTT, R. J. & FORBES, D. 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MOUNTFORD NERC Centre for Ecology and Hydrology, Monks Wood, Abbots Ripon, Huntingdon PE28 2LS ABSTRACT The indigenous distribution of Water Germander Teucrium scordium, an endangered Red Data species in Britain, has contracted to just three sites: two in dune slacks in North Devon and one in a disused flooded limestone quarry in Cambridgeshire. In at least two of these, it has been suffering a decline. At Kingfishers Bridge near Wicken in Cambridgeshire what appears to be a highly successful attempt to boost an ailing indigenous population has been undertaken by transplanting plants propagated vegetatively from the original site onto a nearby wetland created since 1995 on what was intensively farmed arable. The population has _ increased explosively from 1522 shoots on plants introduced in 1997-2000 to over 180,000 shoots in September 2004. The spread appears to be largely by fragile stolon fragments, sometimes broken from the parent plants by grazing geese, being distributed by water in winter and early spring. Optimal conditions for establishment are sparsely vegetated water margins exposed by summer draw-down. The water at Kingfishers Bridge is base-rich and generally of good quality. Water Germander does not thrive in the shade of dense reed, scrub and other tall vegetation, but can tolerate inundation by water and ice cover in winter. Reintroduction also seems to have been successfully achieved at Bassenhally, an indigenous Cambridgeshire site from which Water Germander disappeared soon after its discovery in 1967. Nearly 1700 shoots were counted there in 2004. INTRODUCTION Water Germander Teucrium scordium is a British Red Data plant, classed as endangered in the recently revised Red Data List (Cheffings & Farrell 2005). It is currently restricted as an indigenous species to three sites, though it does occur in Western Ireland. The British populations have not been flourishing in recent years. This paper des- cribes what appears to be a successful attempt to increase the Cambridgeshire population of Water Germander at a wetland recently created from intensive arable farmland at Kingfishers Bridge, close to the only remaining indigenous site in East Anglia and at another Cambridge- shire location from which the species had recently become extinct. This species recovery project is part of a wider ecological restoration campaign in the Fenland basin to rehabilitate degraded wetlands and to create new wetlands where they have been destroyed (Colston 2003; Hughes ef al. 2005; Mountford & Wadsworth 2002). New and rehabilitated wetlands should support, in a sustainable way, as full a range of taxa as possible that were present in Fenland prior to the era of intensive drainage. In the 21st century context of climate change and fragmented landscapes, such restoration cannot hope to create individual perfect facsimiles of previous habitats and species assemblages. However, restoration programmes such as that at Kingfishers Bridge seek to restore natural ecological processes rather than make cons- tructed copies of past habitats and are the antithesis of the “gardening” that has occurred in some nature management (Tomkins 1998). The programme at Kingfishers Bridge has also led to various observations on the ecology of Teucrium scordium which help to confirm the causes of the demise of the species in Britain and also offer ways of boosting existing populations by management and restoring this attractive plant to other sites. 304 R. C. BEECROFT, C. J. CADBURY AND J.O. MOUNTFORD PAST DISTRIBUTION — BRITAIN AND IRELAND Teucrium scordium has never been known to have other than a very localised distribution in Britain. Pre-1970 it had been recorded from 24 10km squares extending from North Devon (v.c. 4), Berkshire (v.c. 22), Oxfordshire (v.c. 23) to South Lincolnshire (v.c. 53), North Lincolnshire (v.c. 54), Southwest Yorkshire (v.c. 63) and Northwest Yorkshire (v.c. 65). The centre of distribution was the East Anglian Fens: Cambridgeshire (v.c. 29) and Hunting- donshire (v.c. 31). By 1960 it had become extinct in all but North Devon, Cambridgeshire and West Suffolk (Perring & Walters 1962, Wigginton 1999, Preston et al. 2002). It was last recorded in South Lincolnshire in 1952 (Gibbons 1975). There is a record from Lakenheath, West Suffolk in 1860 (Hind 1889). It was ‘rediscovered’ in Stallode Wash (Botany Bay), Lakenheath in 1976 by M. G. and S. Rutterford (Rutterford 1977), but had disappeared by 1979 (Simpson 1982) — see Figure 1. In 2004, T. scordium was discovered on Anglesey (v.c. 52) by Richard Birch, where a small patch was found in 2004 in the draw- down zone of a balancing pond. The provenance and status of this population is unknown (Ian Bonner, pers. comm.), and there are no historical records from Anglesey or from Wales. In Cambridgeshire, pre-1860, T. scordium was known from several sites in the fens, including from what is now within the northern boundary of Cambridge City (King’s Hedges and Histon Road), ‘abundantly on the road to Ely’, Roswell Pits, Ely and at Mepal. There were earlier records from Waterbeach and Cottenham which could not be confirmed (Babington 1860). By the publication of the last full Cambridgeshire flora (Perring ef al. 1964), it had long become extinct at all these sites, evidently due to its intolerance of com- petition from tall vegetation. A population does persist in the North Pit, (TL544733) 2:5 km north of Upware on the River Cam and the same distance northwest of Wicken Fen. Moreover, in 1967, Water Germander was discovered by Frank Perring and Ian Hepburn at Bassenhally Pit (TL286986) 1 km northeast of Whittlesey and close to the Nene Washes. By 1970, it had disappeared from this site (Langton 1970), though material from Bassenhally was grown on in the University Botanic Garden, Cambridge. In North Devon, Water Germander has been known since the end of the 18th century from Braunton Burrows (Polwhele 1797). It was locally abundant in the winter-flooded dune slacks well into the 20th century (Martin & Fraser 1939, Palmer 1946). In 1982 a survey by J. Vause, J. Breeds, J. Hope-Simpson and M Tulloh recorded 18,874 shoots at six sites on the Burrows (Vause 1982). In 1998 only 3951 shoots at two of these sites were found. However, four ‘new’ sub-populations contain- ing about 6881 shoots were located, including one of 6604 shoots previously unknown. Excluding the latter, but taking into account all colonies known up to 1997, the 1998 survey revealed 4228 shoots, a 78% decline on the 1982 count (Holyoak 1998). In an attempt to reverse this decline, from 1998 the site managers began to mechanically surface-scrape dune slacks that had formerly supported T. scordium, leading to a substantial population recovery by 2005 (J. Diamond, pers. comm.). A second North Devon location was discovered by Mr W. Tucker in 1976 at Northam Burrows on the opposite side of the Taw Estuary from Braunton, growing in a similar habitat (Ivimey-Cook 1984; Tucker 1978). A survey of Northam made by one of the authors of the present paper (JOM) with Mr Tucker (then North Devon B.S.B.I. recorder) in 1997 showed a small population of <500 shoots, having declined from the few thousand stems of the 1970s, probably due to vegetation succession within the dune slack (W. Tucker, pers. comm.). The North Pit near Upware, Cambridgeshire Braunton Burrows and Northam Burrows currently all have SSSI status. The plant in Devon differs from that in Cambridgeshire in being very hairy and has been described as var. dunense Druce (P. Sell in press). Teucrium scordium has been recorded from twelve 10 km squares in western Ireland since 1970 and since 1987 is known from 10 of these (Preston et al. 2002). It extended along the Shannon from Roosky (Leitrim v.c. H29) to Doonass (Limerick v.c. H8). It is still locally abundant on the calcareous shores of Lough Derg (Clare v.c. H9; and SE Galway v.c. H15) and Lough Ree (southeast Galway). There are isolated stations in North Tipperary, v.c. H10 and elsewhere in Clare (Praeger 1934, Webb OTT): Water Germander occurred on Guernsey in the Channel Islands where it was last seen in 305 TEUCRIUM SCORDIUM IN CAMBRIDGESHIRE L886] VOUIS PIpIOOIY °F L86[ QOUIS popIOdaI UIDq SBY WNIP1OIS “J, dIOYM Sorenbs wry OT °g PopsOdeI Ud9q sey WNIPJOIS “J YOTYM YM sosenbs ury OT [TV “-V ‘SUIMOYS ‘puvloly pue ule Ul "] wnIps1oozs wnisdna] Jo UoNqMSIG “| AMINO Sp1OOOI [TV “VW 306 R. C. BEECROFT, C. J. CADBURY AND J.O. MOUNTFORD TABLE 1. TEUCRIUM SCORDIUM SHOOT COUNTS CAMBRIDGESHIRE SITES 1997-2004 1997 1998 1999 2000 2001 2002 2004 —‘ Total North Pit (original, indigenous) 330 DI 700 1200 558 600 214 (introduced) Ses p; 8 4 3 0 Cam Washes SSSI DoE 1294 156 0 Wetland Creation Project (WCP) Kingfishers Bridge Fen (sites 1,2 & 10) 96* 724110" 1964260" 1000 1500 338 Reedbed (sites 3 & 4) 43% U2 69+800* 042330 2761 IWW25 Lake (sites 7 & 8) S10 86 423 520 3088 =. 41,014 Winter Flood (site 9) 24* 414+114* 165 1970 10,820 134,686 Contour Ditch (site 5) Das 39 119 130 2409 3595 Big Ditch B38 436 Total shoots WCP 310 972 5950) 20 S/S sees Total plants introduced 238% 224* 1060* 0 a3" 1845 Bassenhally Pit 41* 96 30 1683 (Whittlesey) (*Planted introductions) TABLE 2. WATER GERMANDER TEUCRIUM SCORDIUM SHOOT DENSITY AND ASSOCIATED VEGETATION AT KINGFISHERS BRIDGE Teucrium shoot density Associated vegetation height Mean/m? Mean cm Kingfishers Bridge North Pit — indigenous (n = 2) 19 1/7 -OE30:9 Winter Flood N (n = 5) 457 + 540 262-23 S (n =4) 564 +557 21-9) 1056 Winter Flood S (n= 5) 52 +95 DANE? Lake shore draw down (n = 5) 45 +95 sya as 3)93} Lake shore drift line (n = 5) 149 +99 30-8 + 12:0 Reedbed/lagoon margin (n = 2) 368 + 240 DS Oyen) Reedbed — tall & dense (n = 3) 192+116 231-3'= 26-7 Ditches (n = 6) 311342230 Byes) a5 a0) Bassenhally Pit Herb-rich fen 49-4 + 29-2 1925 (McClintock 1975). The Guernsey plant was said to be referable to subsp. scordioides Rouy, which is the commonest form in coastal sites in southern Europe, including the Atlantic fringe and Mediterranean. A recent biometric study of T. scordium populations in northwest France and Brittany suggests that one population in dune slacks in Pas-de-Calais may be referable to subsp. scordioides, with other populations at the site appearing intermediate between subsp. scordioides and subsp. scordium (Toussaint et al. 2005). : BACKGROUND TO THE KINGFISHERS BRIDGE WETLAND CREATION PROJECT Since 1970, attempts have been made to arrest the catastrophic decline of 7. scordium through management of its remaining sites. By 1998, these attempts have met with some local success, but the area of remaining suitable habitat at North Pit (Upware) was tiny, both the Bassenhally and Stallode Wash populations had disappeared and in Devon, T. scordium was either declining (Northam Burrows) or with an TEUCRIUM SCORDIUM IN CAMBRIDGESHIRE 307 uncertain future due to changes in the site’s conservation status (Braunton Burrows). At a regional, national and international scale, it was becoming increasingly clear that nature conservation of small discrete sites through intensive management (‘gardening’) was becoming non-viable (Colston 2003) and that effective protection of nature had to not only conserve that which survived but pursue “creative conservation” i.e. ecological restoration (Sheail er al. 1997). The underlying philosophy behind the Kingfishers Bridge project is precisely this type of creative conservation and seeks to achieve a viable population of T. scordium in a large wetland site that itself forms part of a network of major restored wetlands around the southern margin of the Fenland basin. The project is a private venture to restore a variety of semi-natural fenland habitats on land that was intensively farmed up to 1995. It lies close to the River Cam and is only 2:5 km from Wicken Fen, one of the oldest nature reserves, which is owned and managed by the National Trust. The project is the inspiration of Andrew Green who has dedicated 65 ha of his and his family’s low-lying arable farmland to creating a nature reserve. Roger Beecroft (one of the authors of this paper) has provided major advisory and management input from the start of the project. The construction of the site has been described by Tomkins (1998). Three types of soil are represented on the site: neutral fenland peat, Gault clay of marine origin and Corallian limestone. Most of the water originates as run-off from the land supplemented by some that is pumped from a neighbouring limestone quarry. It is both highly calcareous and of good quality. A small indigenous population of Teucrium scordium grows in a flooded long-abandoned limestone quarry close to the created wetland. Five created habitats suitable for T. scordium at Kingfishers Bridge were well established by 2004: e A 7:0 ha shallow lake of permanent water with eleven islands. In the summer there is considerable draw-down which exposes a sparsely vegetated shoreline; e A 9-4 ha reed-swamp with pools of shallow water and stands of other marsh vegetation (the pools are subject to draw-down in summer); e Seasonally flooded wet grassland in three areas which have shallow pools that remain well into the spring and which are grazed by sheep and wildfowl]; e Highly calcareous margins of two water- filled ditches; e Two quarries specially excavated in Corallian limestone to provide nest-sites for Sand Martins (Riparia riparia) and Common Kingfishers (Alcedo atthis). Each quarry has a pool of clear water at the base of the excavated face. RECORDING METHODS FOR TEUCRIUM SCORDIUM As the species is a stoloniferous perennial, counting plants is impracticable, except for young ones occurring at a low density on bare ground. Following Holyoak (1998), counts of basal shoots made in late August or September were found to be a repeatable methodology allowing comparison of populations at different sites (Table 1). Mean shoot density per square metre was calculated by counting shoots in five 20 x 20 cm quadrats (sub-samples) nested within a 2 x 2 m quadrat. (Table 2). In all but the North Pit (¢ndigenous population) and the Big Ditch (introduced) five 2 x 2 m quadrat samples were taken. The total shoot count was calculated either by counting shoots over the entire habitat (as in most instances) or where densities were particularly high over a sizeable area (as on the Winter Flood) by extrapolation from the mean density /m* x measured area. The quadrat samples were also used to obtain cover/abundance estimates on a Domin scale for Teucrium scordium and associated plant species. Those species occurring in four or more of the sample quadrats in a habitat were treated as constants. One measurement of vegetation height was taken in each of the nested 20 x 20 cm sub-samples within the quadrats (Table 2). Distance from the nearest surface water was also measured as was the depth of ‘soft’ substrate (i.e. penetrable by a narrow pole). Each quadrat was located to a 10 figure grid reference by GPS. POPULATION AND DISTRIBUTION OF WATER GERMANDER AT KINGFISHERS BRIDGE A) INDIGENOUS POPULATION T. scordium has long been known in the North Pit (often referred to as being at Upware though 2:5 km north of the village) at TL54367333. The population has _ been monitored irregularly in terms of percentage cover, but systematic recording (shoot counts) began in 1997 when R. Beecroft recorded 330 shoots. Cutting the Reed and other vegetation 308 R. C. BEECROFT, C. J. CADBURY AND J.O. MOUNTFORD over all the area occupied by T. scordium resulted in an increase to 1200 shoots in 2000. Since then, vegetation cover has increased once more and water levels in the pit have risen. The shoot count diminished to 214 in 2004 and just 35 in 2006. The introduction in 1998 of a few propagated plants to a slightly different area within the pit had failed by 2004 (Table 1). B) PROPAGATION Cuttings from growing shoots were initially taken from the North Pit SSSI in 1997. Cuttings rooted easily with a success rate of almost 100%. The first batch of propagated plants was used as stock to provide more cuttings. Since there was no evident advantage from rooting in water first, subsequent cuttings were placed in compost from the outset. Propagation appeared to be a straightforward process and, once rooted, plants were kept in pots for about a year before being planted out. C) PLANTING OUT Initially 238 propagated plants were introduced to nine sites within the Kingfishers Bridge Wetland Creation Project area on 4th August 1998. Further introductions of propagated plants were made in 1999 (224 plants), 2000 (1060 plants) and 2003 (323 plants) — Table 1. The locations were chosen to give a range of hydrological conditions in an attempt to compare survival under different water regimes. Since it was difficult to identify individual plants once established, progress was measured by counting shoots in August and September. D) INTRODUCTIONS The propagated plants from North Pit stock were introduced to the Kingfishers Bridge Wetland Creation Project area as follows: the Fen, TL545738 (three sites); the Reedbed TL545739 (two sites); the south shore of the Lake TL540737 (two sites); the Winter Flood, TL541737 (one site) and two ditch banks, TL544736 and TL547733. A total of 1845 plants were introduced between 1998 and 2003. From 310 shoots on plants introduced in 1998, the count rose to 5950 by 2001, 20,578 by 2002 and in 2004 the number of shoots had soared to an estimated 181,000. The most marked increases were on the Winter Flood (from 165 shoots in 2000 to an estimated 135,000 in 2004) and the Lake, where plants had spread to all the shores and the seven islands (86 shoots in 2000 to 41,014 in 2004 including 11,060 on the islands). T. ‘scordium has flourished and spread to a lesser degree on - the banks of the two ditches (at one of them, 119 shoots in 2000 to 3595 in 2004). In the Fen and Reedbed, however, there was an increase up to 2002 but a subsequent marked decrease associated with development of a tall and dense reedbed that has invaded the Fen (Table 1). An introduction was made in 2000 to the adjacent winter-flooded Dimmock’s Cote Washes TL541727 (Cam Washes S.S.S.I.) foll- owing slubbing out of the channel and creation of open ground, in order to test establishment under a differing water-regime. The shoot count increased from 599 that year to 1294 in 2001, but by the following year few plants had survived and none was seen in 2004 (Table 1). It appears that the plants had failed to compete with robust Reed Sweet-grass Glyceria maxima. This site also lacks the summer draw- down present in Kingfishers Bridge. HISTORY OF THE POPULATION AT BASSENHALLY PIT (WHITTLESEY) In 1967 a small population (ca 20 plants) of T. scordium was found at Bassenhally Pit on bare mud by a path across a ditch (Crompton 2006). Following its disappearance by 1970, propagated plants from Bassenhally stock were reintroduced to the same location in 1973 by S. M. Walters, R. M. Payne and JOM. The population survived until at least 1975 but became increasingly etiolated and rarely if ever flowered (Crompton 2006; JOM records) it was not subsequently observed. This failure appeared to be due to the particular path having been abandoned and Hawthorn (Crataegus monogyna) scrub having grown up to shade the pit margin. By 1980 the original location appeared entirely unfavourable to Water Germander. A second introduction, this time from North Pit stock, was made in 1999 to a nearby herb-rich largely scrub-free _ fen, TL286987, known as the Central Depression (Langton 1970). Here where there are patches of relatively short vegetation, Water Germander had done well, and the count of 41 shoots introduced in 1999 had increased to 1683 by 2004 (Table 1). HABITAT I. KINGFISHERS BRIDGE AND UPWARE NORTH PIT a) North Pit The flooded North Pit, a long abandoned Corallian limestone quarry, supports the indigenous 7, scordium population, which has been declining since 2000, possibly as a result of shading by taller vegetation and scrub that has invaded the pit, and variation in water quality and depth. TEUCRIUM SCORDIUM IN CAMBRIDGESHIRE 309 The vegetation of two 2 x 2 m quadrats in which T. scordium grows has been surveyed annually 1997-2004 with the exception of 2003. The NVC community of the site is Reed swamp (S4: Rodwell 1991-2000), and specifically the Marsh Bedstraw (Galium palustre) sub-community (S4b). Constants throughout the period have been Phragmites australis, Galium palustre, Great Pond Sedge (Carex riparia), Great Reedmace (Typha latifolia), Lesser Duckweed (Lemna minor) and the moss Drepanocladus aduncus. Lesser Water-parsnip (Berula erecta) and Great Water Dock (Rumex hydrolapathum) were constants in most years. Comparison with a survey made in 1989 (JOM and G. Crompton) suggests that Ivy-leaved Duckweed (Lemna trisulca) and Bittersweet (Solanum dulcamara) have both declined, the former markedly so. The overhanging scrub is composed mainly of Grey Sallow (Salix cinerea), Hawthorn (Crataegus monogyna) and Guelder-Rose (Viburnum opulus). In 2004 the reed stems were ‘spindly’ and growing at a rather low density, with a mean height of 177-0 + 30-9 cm. The Reed swamp was floating and at the site where T. scordium was growing the water depth at the surface was 5—-10cm with 30—40cm of silt below. T. scordium did not extend into the wetter and increasingly quaking swamp in the centre of the pit. b) Reedbed and Fen The Reedbed was established from what had been peaty arable land in 1997. T. scordium was introduced into the young reedbed first in 1998 and again in 2000. By 2004, a dense S4 reed-swamp had developed with Phragmites of an average height (in September 2004) of 231:3 cm (n = 15). There was little other vegetation in the reedbed apart from 7. scordium, which struggled to compete. Etiolated plants of T. scordium reached 180 cm in height, but had few if any flowers. The Reedbed bordered a shallow lagoon, which by the end of the summer had an extensive draw-down zone of peaty mud colonised by Water Mint (Mentha aquatica), Amphibious Bistort (Persicaria amphibia), Creeping Bent (Agrostis stolonifera) and the moss Drepanocladus aduncus as well as sparse Phragmites. In 2004 T. scordium was flourishing in this open draw-down area between the reedbed and lagoon. The mean density there was 308 + 240 shoots/m? compared with 192 + 116 shoots/m* in the dense reedbed. Virtually all the Reedbed apart from embankments is submerged in shallow water over the winter. The Fen, on somewhat higher ground, had a vegetation community that was probably best described as 826 tall herb fen Phragmites australis—Urtica dioica (Rodwell 1991-2000). Much of it has however converted to reed-swamp. c) Winter Flood This is an area of grassland bordering the Lake. Its depressions are shallowly flooded for much of the winter and spring. With a depth of only 5 cm of peat above the clay substrate, drainage was poor. In the northeast sector where T. scordium was particularly abundant, the constant plant species (forming over 50% cover in each sample) were Persicaria amphibia, Mentha aquatica, Whorled Mint (Mentha x verticillata), Gypsywort (Lycopus europaeus), Scentless Mayweed (Tripleurospermum inodorum), Great Plantain (Plantago major) and Agrostis stolonifera. Over an area of 0°13 ha T. scordium was one of the dominant species with an average density of 564 + 557 shoots/m’. However, where Mentha aquatica formed the predominant vegetation, T. scordium was sparse. Further from the Lake, 150 m south of the main population, 7. scordium was growing around depressions in the grassland that were permanently inundated between November and April/May, but only intermittently flooded by rain water at other times. Here the constants were Agrostis stolonifera, Common Spike-rush (Eleocharis palustris), Jointed Rush (Juncus articulatus) and Plantago major. The density of T, scordium was 52 + 95 shoots/m’, less than a tenth of that of the main population. Such vegetation is not unequivocally classified in the NVC, but shows some allegiance to various kinds of regularly inundated grasslands within the alliance Elymo-Rumicion crispi e.g. MG11 Festuca rubra-Agrostis stolonifera-Potentilla anserina grassland, MG13 Agrostis stolonifera- Alopecurus geniculatus grassland and OV28 Agrostis stolonifera-Ranunculus repens community (Rodwell 1991-2000). d) The Lake At its full extent there are 7 ha of water with eleven islands, some of which are submerged in winter. In summer a considerable area of peat and clay is exposed on the Lake shore and around the islands by draw-down. Such areas, 310 R. C. BEECROFT, C. J. CADBURY AND J.O. MOUNTFORD including all the islands, have been extensively colonised by 7. scordium. The main constants in the draw-down zone on the south shore were Mentha aquatica, Redshank (Persicaria maculosa), Hemp Agrimony (Eupatorium cannabinum), Brookweed (Samolus valerandi), Toad Rush (Juncus bufonius) and Lycopus europaeus. A number of annuals such as Red Goosefoot (Chenopodium rubrum) and Golden Dock (Rumex maritimus) were frequent. The average density of 7. scordium was 45 + 95 shoots/m* (n = 25), with 50-90% of the draw- down zone still bare peat. Many of the plants had only become established for a year or two. At the winter/spring high water mark on the southern shore of the lake there was a 2 m wide drift line. This was totally vegetated, predominantly by Lycopus europaeus, Mentha aquatica, Eupatorium cannabinum, Agrostis stolonifera, Persicaria amphibia, Samolus valerandi and such annuals as Persicaria maculosa and Trifid Bur-marigold (Bidens tripartita). T. scordium here attained a mean density of 149 + 99 shoots/m* (= 25): e) Ditches T. scordium was introduced at two sites on the margins of ditches (Big Ditch and Contour Ditch). Here the substrate was _ highly calcareous clay. Where Teucrium was growing, the constants were Agrostis stolonifera, Samolus. valerandi, Plantago major and Eleocharis palustris. The vegetation height in the sample plots varied between 11:4 cm and 78:.8 cm (n = 7). The T. scordium density was highly variable ranging from 35 to 520 shoots/ 0b (G0 7). II. BASSENHALLY PIT Here the 7. scordium occupied two distinct vegetation communities when surveyed in 2004. Where it had been reintroduced in 1999, Agrostis stolonifera, Brown Sedge (Carex disticha), Slender Spike-rush (Eleocharis uniglumis), Galium palustre and the moss Drepanocladus aduncus were dominant and the community was again best placed within the Elymo-Rumicion crispi. The presence in the site of Eleocharis uniglumis, Juncus gerardii and, in the second vegetation community, Schoenoplectus tabernaemontani suggested some brackish influence, and relationship to SM20 Eleocharis uniglumis — saltmarsh (Rodwell 1991-2000). The physiognomy of the vegetation has some resemblance to a dune- slack, and some areas resembled SD17 Potentilla anserina-Carex nigra dune-slack, another component of the Elymo-Rumicion crispi alliance that is normally coastal in the UK (Rodwell 1991-2000). 329 shoots of T. scordium were counted in this zone. T. scordium had also spread into taller vegetation in which Slender Tufted-sedge (Carex acuta) was predominant and included such species as Marsh Pennywort (Hydrocotyle vulgaris) and Grey Club-rush (Schoenoplectus tabernaemontani). Eighty percent of the population of T. scordium (1683 shoots) occupied this community which may prove to be a variant of S6 Carex riparia swamp (Rodwell 1991-2000). The mean shoot height of T. scordium at Bassenhally was 38-7 + 19-4 cm, while that of the surrounding vegetation was 49-4 + 29:2 cm. The Central Depression at Bassenhally Pit is the most important part of a - site with many species typical of herb-rich fen, including some locally scarce plants such as Great Water-parsnip (Sium latifolium), Marsh Speedwell (Veronica scutellata), Lesser Water- plantain (Baldellia ranunculoides), as well as Eleocharis uniglumis. REPRODUCTION AND SPREAD: KINGFISHERS BRIDGE T. scordium flowered prolifically in the more open areas at Kingfishers Bridge (Lake Shore, Winter Flood, the draw-down zone at the edge of the Reedbed and ditch margins). In the dense reedbed and in the North Pit where plants were shaded, flowering was largely suppressed. Many plants that had flowered on the Winter Flood were examined on 12 October 2004 — though abundant seed had been set with two to four nutlets per calyx, very few seeds had ripened by that date. At the time of the survey in mid-September 2004, there was little evidence of reproduction from seed, even on the bare peat of the Lake shore. Most reproduction, at least at Kingfishers Bridge, appears to be vegetative from stolons that are produced in late summer and autumn. They differ from the erect flowering shoots in extending horizontally from the plant and are thickened, suggesting a storage function. In winter, stolon segments get detached from the parent plants e.g. by trampling and grazing by wildfowl — particularly Greylag Geese (Anser anser) and Canada Geese (Branta canadensis). Though sheep may trample and fragment T. scordium by trampling, they appear not to eat it, possibly because of its garlic-like odour". ‘Both German vernacular names (Knoblauch-Gamaner and. Lauch-Gamander) and the specific epithet scordium itself refer to garlic or leek. TEUCRIUM SCORDIUM IN CAMBRIDGESHIRE 311 Stolon fragments have been seen floating in flood water, even under ice. 7. scordium had spread naturally from the original planting of 50 plants on the south central shore of the lake in 1998 right round the margin and to all eleven islands by 2004. In addition, a new colony appeared naturally on the east side of the Big Ditch opposite a planting on the west bank, 15 m across the water. On the Winter Flood even low raised banks 20-30 cm high have withheld the flow of winter flood water across the area and this has apparently limited the spread of TJ. scordium westward and southward. However, by 2005, plants had become established on a nearby, but discrete, winter-spring flooded area, in the Wader Meadow. This colonisation probably originated as a result of water being pumped onto the site from the Lake. By September 2006, there had been a further massive increase in the 7. scordium cover, both around the Lake and on the sheep — and goose-grazed Winter Flood, where over an area of 0:58 ha it was dominant almost to the exclusion of other plants. PRELIMINARY CONCLUSIONS ON THE ECOLOGY OF WATER GERMANDER IN CAMBRIDGESHIRE The introduction of 7. scordium has revealed or confirmed certain aspects in the ecology of the species (Hill et al. 1999): e it flourishes in calcareous substrate — clays and peat (and calcareous sand in North Devon dunes) — the Ellenberg R indicator value is 8 i.e. moderately basic; e it requires damp conditions and tolerates long periods of submergence in winter but quaking swamp conditions through the summer are detrimental — the Ellenberg F indicator value is 8 1.e. sites that are damp to seasonally wet; e flowering is suppressed by shading, and though shoots become etiolated it appears that perennial plants are eventually out- competed by tall dense vegetation (such as Reed and scrub) and fail to reproduce vegetatively — the Ellenberg L indicator value is 7 i.e. preferring well-lit places, but will survive for a while in partial shade; e the main form of reproduction in the studied sites is vegetative by stolons, which when detached can be dispersed widely by water; e low banks that inhibit the flow of water can act as barriers to vegetative dispersal; e optimal conditions for colonisation are the sparsely vegetated sites that are exposed on the margins of water bodies by summer draw-down. ECOLOGY OF TEUCRIUM SCORDIUM IN NORTH DEVON Most of the colonies of Water Germander in North Devon are in dune slacks that have, at least until recently, been regularly inundated by a water-table rise in winter. Inundation may continue well into the spring. Two of the sites were margins of artificial ponds. The large colonies were on predominantly level ground. The substrate on which Teucrium was growing was calcareous sand that was fairly humic (Holyoak 1998). Holyoak emphasised that Water Germander on the Burrows was assoc- iated with short vegetation, much of it less than 5 cm, and grew sparsely where vegetation exceeded 15 cm. It colonised temporarily disturbed ground after the excavation of ponds, and could tolerate some vehicle pressure leading to ruts. The short vegetation was being maintained by rabbit grazing and the poor nutrient status of the soil, together with winter flooding and _ near-drought conditions in summer. A large number of plant species were associated with Water Germander in the Devon dune slacks. Most constant associates were Agrostis stolonifera, Bog Pimpernel (Anagallis tenella), Marsh Helleborine (Epipactis palustris), Hydrocotyle vulgaris, Sharp Rush (Juncus acutus), Sea Rush (Juncus maritimus), Mentha aquatica, Silverweed (Potentilla anserina), Common Fleabane (Pulicaria dysenterica) Creeping Willow (Salix repens) and the moss Drepanocladus aduncus (Holyoak 1998; unpublished data JOM). Such vegetation again approaches NVC communities of the Elymo-Rumicion crispi such as MG11 and SD17, as well as other dune-slack types that are either more fenny e.g. SDI5 Salix repens—Calliergon cuspidatum or more scrub- dominated e.g. SD16 Salix repens-Holcus lanatus (Rodwell 1991—2000). Features similar to the Kingfishers Bridge site were: | e winter-spring flooding and tolerance of inundation; e calcareous substrate with some shallow humic soil; e short vegetation; 312 R. C. BEECROFT, C. J. CADBURY AND J. O. MOUNTFORD e colonisation of temporary disturbed ground; and e frequent associated plants in all the Devon and Cambridgeshire sites were Agrostis stolonifera, Mentha aquatica and Drepan- ocladus aduncus. Hydrocotyle vulgaris occurred at Bassenhally and in the Devon slacks. Major differences were: e Braunton Burrows and, to a lesser extent, Northam Burrows are essentially natural; the Kingfishers Bridge wetland is recently created, even where the indigenous popu- lation of 7. scordium still occurs is a disused quarry. Bassenhally Pit also is artificial, though like the North Pit, it has been abandoned for many decades; e in Devon the main habitat is dune slack (to which the Bassenhally site bears some resemblance); at Kingfishers Bridge the species has flourished in the draw-down zone around a lake, reedbed lagoons and seasonally flooded grassland resulting from lowering water-table in summer; e the substrate in Devon is humic sand; at the Fenland sites it is clay with some peat; e on the Burrows _ species frequently associated with 7. scordium included Epipactis palustris, Potentilla anserina, Pulicaria dysenterica and Salix repens, all of which were absent from sites where Water Germander occurred at Kingfishers Bridge. There Persicaria amphibia, Lycopus europaeus, Plantago major and Samolus valerandi were among the most frequent associates. A mixture of Carex species and submaritime Cyperaceae were especially common at Bassenhally; e the areas where T. scordium still flourished at Braunton and Northam were heavily grazed by rabbits, which also browsed the margins of the Bassenhally site, while those at Kingfishers Bridge were grazed by sheep and feral geese. REASONS FOR THE DECLINE IN THE UK It should be acknowledged that, as a species of early successional habitats, T. scordium may always have been transient in individual sites, but able to maintain itself in a complex of sites within a region since, as one site became overgrown, suitable conditions developed else- where nearby. Insofar as the available literature . can be used to reconstruct its distribution through the past 250 years, there is evidence that 7. scordium indeed survived for many decades in particular regions, though “monitor- ing data” for particular populations were only gathered from the latter part of the 20th century onward. The main reasons for the disappearance of Water Germander from most of its British sites were probably drainage and land claim. However, within those sites that survived into the modern era, it appears that competition and shading from tall wetland vegetation are the main factors in its decline (Perring et al. 1964). Thus at Stallode Wash at Lakenheath, its demise was almost certainly as a result of being unable to compete with tall vigorous Phrag- mites australis (Wigginton 1999). Indeed surveys made of the former Lakenheath site in the 1990s showed almost 100% cover of dense Reed, with canes reaching >2°5 m tall (JOM unpublished data). Its decline and lack of vigour at North Pit, Upware appears to be due to shading by scrub, though higher water levels may have contributed leading to extensive quaking swamp. Poorer water quality may also be a factor, with turbid water during winter floods preventing growth — witness the con- temporary decline of Lemna trisulca at Upware. The fact that the indigenous population at Bassenhally Pit died out and that the first reintroduction was unsuccessful was almost certainly also” ‘duc tomeserub encroachment and the end of light trampling at that particular location within the pit. Both in Britain and Ireland, the most vigorous T. scordium populations have occurred where there is light disturbance, through trampling, peat cutting or regular draw-down. In the Reedbed at Kingfishers Bridge 7. scordium still survives, but is unlikely to persist for long in the dense vegetation. The failure of the introduction at Dimmock’s Cote is also linked to a highly competitive grass (Glyceria maxima). At Braunton Burrows the decline is attributable to a falling water table and scrub encroachment in the dune slacks. Holyoak (1998) states that the vegetation had grown markedly taller between the 1982 and 1998 surveys. At a few sites where Water Germander was still flourishing the vegetation was kept short by rabbit grazing. Remedial conservation action at Braunton in 1998 has met with early success, and by 2004 T. scordium had re-appeared in seven of the eight dune-slacks where the surface has been TEUCRIUM SCORDIUM IN CAMBRIDGESHIRE 333 mechanically scraped (J. Diamond, pers. comm.). This recovery appears to be due both to reduced competition from Salix repens, Juncus acutus etc., and also to bringing the sand surface where 7. scordium is rooted closer to the underlying water-table. Attention must also focus on the regeneration niche for 7. scordium, and whether the species reproduces primarily from _ seed, or vegetatively. Data on seed productivity and viability come from collections of T. scordium held in the Royal Botanic Gardens Kew’s Millennium Seed Bank at Wakehurst Place. From this source it is clear that 7. scordium may produce very little viable seed in the wild, with collections from British populations varying from a maximum of 40 viable seeds per plant (i.e. 40 plants producing around 1600 seeds) to as little as 1 viable seed per plant (i.e. 12 plants producing an overall total of just 12 viable seeds). When wild material is grown on in horticultural conditions, productivity may be much higher, with the total seed-output of 40 plants varying from 1350 to ca 51000 seeds. A very small trail conducted within the present study is consistent with these results. Seeds collected from North Pit were sown to check viability, and only one seed germinated from twelve sown in March onto saturated compost and kept in a poly-tunnel. Taken together these observations strongly suggest that vegetative reproduction will be more important than establishment from seed in wild British populations; and may provide further explan- ation of why T. scordium has not re-established when previous sites have been subject to apparently suitable restoration management. THE CAMBRIDGESHIRE SITES IN A EUROPEAN CONTEXT According to Clapham eft al. (1987), T. scordium is confined to a) Europe from Sweden, notably Oland and Gotland (Hultén 1950) and Estonia southwards and b) to West Siberia and the Aral-Caspian region. Hultén and Fries (1986) essentially confirm this distribution, showing it to be uncommon throughout its range but with concentrations locally in Germany, the Caucasus and former Yugoslavia. Hultén and Fries also suggest that T. scordium subsp. scordioides is largely confined to the Mediterranean zone, the Black Sea and Caucasus and the Aral-Caspian, although other sources (e.g. McClintock 1975) indicate that this subspecies is found as far north as the English Channel. The southern limits of its native distribution appear to be in Morocco (seemingly absent elsewhere in Africa), Malta (where it is endangered) Lebanon, Israel, northern Iran and Uzbekistan. Recent national distribution maps for countries in Central Europe indicate that 7. scordium is most commonly associated with the floodplains of major rivers, and there is a suggestion that it has declined sharply in the former Federal Republic of Germany and Switzerland, whilst remaining locally frequent in the former Democratic Republic (Haeupler & Schénfelder 1989; Lauber & Wagner 2002; Benkert et al. 1998). Elsewhere in central and northern Europe, 7. scordium is threatened in the Czech Republic, and endangered in both Latvia and Lithuania. Though in Britain, T. scordium is at the edge of its distribution in Europe, some insight into its ecological requirements and causes of decline can be inferred from its behaviour elsewhere. In Ireland and the Netherlands, it is a typical species of the phytosociological alliance Elymo-Rumicion crispi (Nordhagen 1940), occurring in flooded swards (Westhoff & den Held 1969; White & Doyle 1982). Such vegetation is typical of seasonally inundated situations (including dune-slacks, lake banks and depressions where rain-water collects) with a marked summer to autumn draw-down leaving expanses of moist habitat with relatively low cover, and hence little com- petition. Although widespread in Europe, typically such vegetation occurs in more humid climates and where the site is liable to disturbance or intensive grazing, suppressing shade and maintaining open soil areas. The Elymo-Rumicion. is_ classified by EUNIS/ CORINE as 37-24 Flood Swards and Related Communities. Agrostis stolonifera is_ the characteristic grass in such communities (Ellenberg 1988). Examination of the SOPHY data base shows that in France T. scordium is mostly lowland or coastal and that amongst its usual associates (or species with markedly similar ecological requirements) are: Eleocharis palustris, Purple Loosestrife (Lythrum salicaria), Mentha aquatica, Persicaria amphibia, Potentilla anserina and — Sium latifolium. Several of these are important species in the Cambridgeshire sites for T. scordium. Interestingly, two other species cited by SOPHY as having a similar ecology to 7. scordium are Ribbon-leaved Water-plantain (Alisma gramineum) and Fen Ragwort (Senecio 314 R. C. BEECROFT, C. J. CADBURY AND J. O. MOUNTFORD paludosus), both extremely rare in Britain, typical of draw-down habitats, and with all or most of their recent sites within the Fenland basin. Within central Europe, 7. scordium appar- ently occurs in more closed fen habitats dominated by large sedges (Wagnocaricion) or in fen-meadows of the Molinietalia (Oberdorfer 1977-1983). Both these vegetation groups are often associated with the Elymo- Rumicion at the same sites 1.e. the large sedge communities occurring where disturbance/ grazing is reduced and the fen-meadow where flooding is less prolonged and regular, and where mowing is often combined with grazing to suppress shrub-encroachment. Further south in Europe, T. s. scordium is often replaced by T. s. scordioides, though both subspecies occur in Romania and France, for example. Within this region, Water Germander is still largely confined to seasonally wet habitats, often on somewhat calcareous sands in the coastal zone e.g. in Greece and Romania (JOM unpublished data). Many of the associated species in such habitats are once again draw-down specialists, including several that are extremely rare in the UK e.g. Brown Galingale (Cyperus fuscus) and _ Jersey Cudweed (Gnaphalium luteo-album), as well as widespread species like Samolus valerandi and dominant Agrostis stolonifera. In some sites near its southern limit of distribution, T. s. scordioides is found in more closed vegetation e.g. Mediterranean tall humid grasslands of the Molinio-Holoschoenion. CONSERVATION OF TEUCRIUM SCORDIUM IN BRITAIN When the success of the Kingfishers Bridge project is assessed in the context of what is known of the ecology of Water Germander, it is clear that the Fenland sites where T. scordium survived into the 20th century were in many respects becoming unsuitable. The evidence from the literature is that these habitats were suitable until possibly the middle of the century, when the reduced disturbance and grazing of wetlands, coupled to drainage and regulation of water-levels resulted in the relict sites becoming overgrown with shrubs and tall reeds (Crompton 2006; Perring et al. 1964). What evidence can be marshalled from the Devon sites suggests that falling water- tables and scrub encroachment were again major factors. Those healthy populations of this species observed by JOM in Ireland and . mainland Europe are frequently in early successional habitats that share natural summer draw-down with low competition, often faci- litated by grazing in lowland calcareous situations. It is apparent that the managers at Kingfishers Bridge have produced precisely these conditions in the early years of the wetland creation project. The continued success of the T. scordium introduction scheme at Kingfishers Bridge requires that these conditions be maintained through periodically renewed disturbance. Changes in T. scordium abundance in the reed swamp at Kingfishers Bridge and recent history at Bassenhally and Braunton strongly suggests that intervention is necessary to prevent succession eliminating suitable conditions. So long as T. scordium and similar draw-down specialists are confined to a few British sites, some interventionist manage- ment may be the only way to assure their survival, though the Kingfishers Bridge project offers the possibility of engineering new sites and introduction of propagules as a practical means to widely disperse these plants through the landscape. However, creation of several large restored wetlands in proximity to one another offers a more sustainable future. Kingfishers Bridge forms part of a new network of wetlands in Fenland that should be sufficiently large and intrinsically diverse for natural ecosystem processes. to ensure that early successional stages are always rep- resented somewhere within a dynamic range of habitats. There is evidence that 7. scordium occurs in rather different habitats in different parts of its range, possibly reflecting genetic differences. Biometric studies of the British and Irish populations, similar to those conducted by Toussaint et al. (2005), might help tailor management prescriptions to the needs of the local populations. There is concern that both the re-introduced Cambridgeshire populations derived from vegetative propagation will have low genetic diversity. Further research is thus required on the relative contributions of sexual and vegetative reproduction (including seed dormancy) in British, Irish and Continental populations of 7. scordium, and the precise conditions required for seed germination and seedling establishment. However, results from the Millennium Seed Bank (R.B.G. Kew at Wakehurst Place) suggest that establishment from seed is much less important that vegetative reproduction in Britain. Evidence TEUCRIUM SCORDIUM IN CAMBRIDGESHIRE Byil5) from site management in the 1990s at both North Pit (Upware) and Bassenhally suggests that 7. scordium does not reappear from dormant seed if suitable conditions for its growth are re-established. Our work with T. scordium together with the 6 wetland restor- ation schemes in Fenland offers realistic hope for building a sustainable mosaic of wetland habitats in Fenland, but still leaves some questions unanswered. ACKNOWLEDGMENTS First and foremost we would like to congratulate Andrew Green for his foresight and pioneering efforts to create a_ highly successful wetland nature reserve at King- fishers Bridge. This has_ received strong Support and encouragement from his family and the Kingfishers Bridge Wetland Creation Trust. It has been a fascinating experience seeing the development of the various habitats on the reserve and their fauna and flora. Andrew Green, Bruce Martin and John Oates were of major assistance in the task of counting Water Germander shoots in September 2004, as was Bill Meek (C.E.H.) at Bassenhally. We are also very grateful to Phil Grey (Whittlesey Wildfowlers — affiliated to BASC) for arranging permission to visit their site at Bassenhally Pit. David Holyoak kindly allowed us to refer to his report to English Nature on Water Germander at Braunton Burrows, North Devon. Bill Tucker and Bob Hodgson, successively B.S.B.I. recorders for North Devon, guided us to the Devon sites and provided historical information. Very helpful information on recent successes in Devon was provided by James Diamond (Natural England) and by Ian Bonner des- cribing the exciting discovery of this plant on Anglesey. John Dickie, R.B.G. Kew provided useful information derived from databases within the Millennium Seed Bank. The propagation of T. scordium, its introduction to the Kingfishers Bridge wetland and subsequent surveillance received very welcome financial support from English Nature’s Species Recovery Programme, and much practical help with the propagation phase provided by Mike Crewe. The maps of Teucrium scordium (Figure 1) were prepared by Henry Arnold (Biological Records Centre, C.E.H. Monks Wood). REFERENCES BABINGTON, C. C. (1860). Flora of Cambridgeshire. J van Voorst, London. BEECROFT, R. (1999). Population recovery of Teucrium scordium at Upware Pit SSSI. 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Joint Nature Conservation Committee, Peterborough. (Accepted October 2006) Watsonia 26: 317-325 (2007) 317 The Eleocharis mamillata H. Lindb. fil. aggregate (Cyperaceae) in the British Isles G. A. SWAN $1, Wansdyke, Morpeth, Northumberland NE61 3QY and A. J. RICHARDS School of Biology, Ridley Building, University of Newcastle NEI 7RU ABSTRACT We analysed stylopodium characters, perianth bristle number, vascular bundle number, stomatal length and fruit density for 50 stems of Eleocharis mamillata agg. throughout its British range. Stylopodium shape varies considerably within a spikelet and it is necessary to measure a large sample of fruits to establish an accurate reading. However there is highly significant variation between individuals for stylopodium shape and _ relative stylopodium width. Variation in these attributes between individuals is continuous. Nine individuals sampled, eight from N.W. Yorks (v.c. 64) and one from Selkirk (v.c. 79), have a stylopodium shape considered typical of subsp. mamillata, which has not been previously recorded from the British Isles. However, this material does not correspond to subsp. mamillata in any other character. Most individuals from the Borders region (v.cc. 67 and 70) are clearly referable to subsp. austriaca (Hayek) Strandhede, but many individuals in N.W. Yorks, and one from Selkirk belong to a distinctive intermediate type which is also distinguished by a high fruit density and vascular bundle number. We hypothesise that these may represent hybrids between previously isolated populations which have been recently transported into water bodies by ducks. KEYWORDS: stylopodium, vascular bundle, hybrid vigour, Spike-rush. INTRODUCTION Eleocharis mamillata H. Lindb. fil. and its close relatives including FE. austriaca Hayek are distinguished from E. palustris (L.) Roemer & Schultes in having fragile stems with an orbicular rather than oval cross-section, fewer vascular bundles, conical flowering heads, and the number of perianth bristles at the base of the fruit usually exceeding four (Walters 1980). The E. mamillata aggregate are plants of a boreal or montane distribution and ecology in Europe, rarely inhabiting the lowland, more eutrophic locations that are typical of E. palustris. E. austriaca was first collected in the British Isles in July 1947 by N.Y. Sandwith in a marshy ox-bow of the infant R. Wharfe below Buckden, Mid-W. Yorks. (v.c. 64) but was not identified until 1960 (Walters 1963). Shortly afterwards, it was discovered in similar habitats by streamsides in montane, usually forested areas, in S. Northumberland (v.c. 67), Cumberland (v.c. 70) (Walters 1963, Swan 1993, Halliday 1997), in Selkirk (v.c. 79) (Corner 1975) and in a number of further localities in the valleys of the R. Wharfe and R. Ribble in v.c. 64 (Roberts 1977). The exact habitats favoured by this’ species are impermanent since several are artificial water bodies and all are colonised by seral dev- elopment resulting in conditions which become unsuitable to this primary coloniser. In v.c. 67 at least, the species has become very rare in recent years. Although it has been traditional to separate E. austriaca from E. mamillata (Hayek 1910, Hegi 1966, Hess et al. 1967, Walters 1980), the two taxa are very similar in general appearance and are clearly closely related to one another. The character by which they are most readily separated is the shape of the stylopodium (the small discrete organ which links the top of the fruit to the style base) which all authors agree is wider than long in E. mamillata and longer. than wide in E. austriaca. The perianth bristle number is said by Hayek to be more commonly Six or more in the former species, but five in the latter, although neither Strandhede (1966) nor Walters (1953) confirm this. E. mamillata 318 G. A. SWAN & A. J. RICHARDS has also been said by Hayek to have, on average, fewer vascular bundles in the stem (not exceeding 12). These are plants with very few quantifiable characters, so the possibility should be borne in mind that any character that shows variability, however trivial, had been given undue importance. This may _ be particularly true of stylopodium characters which we have found can vary greatly depending on the position of the fruit in the spike. It has been suggested that FE. austriaca occurs in mountainous, calcareous areas, and is localised to parts of Norway within Scandinavia, whereas E. mamillata is more typical of peaty pools without competing vegetation and is more widespread in Scandinavia, although both species are common in central Europe and the Alps (Strandhede & Dahlgren 1968). However, Strandhede (1966) recombined E. austriaca as E. mamillata H. Lindb. fil. subsp. austriaca (Hayek) Strandhede and recently he has been followed by others, especially Gregor (2003), who provided considerable biometric support for this treatment; it is evident, therefore, that the correct name for the British plant at specific rank should be E. mamillata. All the material that has been examined previously from v.cc. 67 and 70 had a long and narrow stylopodium and were presumably correctly referred to subsp. austriaca (E. austriaca). However, fruits from a Yorkshire population in Strandhede (1966; see Fig. 7e) appear to have characters intermediate between austriaca and mamillata whilst drawings of some fruits in Walters (1963) and Roberts (1977) also led Gregor (2003) to consider plants from northern England to be typical of neither subspecies. Furthermore, some material sent to G. A. Swan (GAS) from Ribbledale and Wharfedale by FJ. Roberts (FJR) was characterised by a broader and wider stylop- odium. At the time GAS considered that this material fell within the limits of subsp. mamillata, which had not been recorded from the British Isles. For many years, GAS had been aware that the fruits of the plant drawn by Roberts (1977) differed from those of the plant which he (GAS) named subsp. austriaca. However, it was only when A. J. Richards (AJR) sent to GAS a copy of Gregor (2003) that GAS felt compelled to try and clarify the situation and he is grateful to AJR and FJR for catalysing and promoting the (albeit incon- clusive) work published here. MATERIALS AND METHODS Growth in EF. mamillata agg. is such that it is impossible to satisfactorily distinguish the limits of clonal individuals in the field. Populations typically have a rather even distribution of shoots that may represent many individuals that overlap one another, or may consist of only one or very few individuals. A total of 50 shoots were examined from 24 populations from various parts of the British range (Table 1). Voucher specimens of these are to be lodged at the Hancock Museum, Newcastle upon Tyne (HAMU). Most populations are represented by only a single specimen, but where more than one specimen was examined, it is not known for certain Whether more than one _ individual was involved, although measurements suggest that this was the case. In order to eliminate bias due to the position of the fruit within the spike, all fruits in a spike were sampled as far as this was possible, and their position in the spike noted. 2300 fruits were analysed in total. Often, some fruits were missing or had dehisced, but it is doubtful if these absences seriously affected the data. In order to count perianth bristle number, fruits need to be mounted in water on a microscope slide but it did not prove possible to accurately count this in some specimens as the bristles on many fruits had been broken or lost. Dry fruits were measured by means of a calibrated eye-piece scale at x20 using a Swift dissecting microscope. For each fruit, the following characters were noted: e Ratio of the length to the maximum width of the stylopodium, known as ‘stylopodium shape’ (s.s.). (The length of the stylo- podium is unambiguous in material which has lost style branches, although the style may not always break in the same position, causing a possible source of inaccuracy; in the few cases where style branches were still intact, the end of the stylopodium was judged to be where the style base reached its minimum width, which probably accords with the definition of Strandhede (1966), although this position is not always assessed readily). e Ratio of the maximum width of the stylopodium to the maximum width of the fruit, known as ‘relative stylopodium width’ (r.s.w.). ELEOCHARIS MAMILLATA sll) TABLE 1. DETAILS OF SHOOTS EXAMINED. SHOOTS WERE CONSIDERED TO BELONG TO THE SAME POPULATION WHEN THEY OCCURRED IN THE SAME WATER BODY WITHIN ABOUT 100 M Code No. of V.C. Site name and code Grid Ref. So iSowee Date Collector fruits 5m 39 64 Lodge Pond, Settle 5 SD822621 0:912 0496 28/09/75 FIR 31m 36 64 Ribble 12 SD777790 =—-0:898 )—_-(0:-496 = 28/08/75. —~FJR 32 m DS 64 Ribble 7 SD782786 0-948 0-459 15/09/03 FJR 40 m 67 64 ~ Ribbledale SLQ 8 SDITS785 0-973, (0445) 12/07/03) VEIR 51m 35 64 Ribblehead SLQ 8 SD773784 0-885 0-429 12/07/03 FJR 53 m 27 64 Ribblehead SLQ 8 SD773784 0-881 0-445 12/07/03 FJR 42m 26 64 Ribble 21 SD785769" ~~ 0:957 0-455), 15/09/03" “FIR 30m 2 64 Buckden, Wharfe 11 SD944760 0-925 0-482 11/09/03 FJR 8m DD, 79 Tima Water ?pop NT20 0:960 0-445 06/07/77 OM Stewart el 53 64 Ribblehead Quarry 3 SD7678 1:156 0-414 04/09/04 FJR on 45 64 Ribble 7 SD782783 = 1:084 0-436 15/09/03 FJR 141 Il 64 Ribble 7 SD784780 1:100 0-418 15/09/03 FJR 101 84 64 Ribblehead SLQ 8 SD773785 =1:073. 0-426 =26/08/75_ FIR 39) il 74 64 Ribbledale SLQ 8 SD773785 =1:043 0-442 12/07/03 FJR jiikat TY 64 Ribble 9 SDI/S57718> 1-018 0-436) 15/09/03 FIR Dai 36 64 = Selside, Ribble 10 SD788787 = 1:017 0-419 18/10/73 FJR Ali WT 64 Ribble 12 SD778789 1:049 0-407 03/09/03 FJR 381 WS 64 Ribble 20 SD798745 =-1:156 =0-417_—-15/09/03_-FJR AS i 41 64 Horton, Ribble 22 SD79.74. 1:068 0-428 14/07/75 FJR Si i 28 64 BenRhydding gravel pit 24 SE146475 =1:086 =0-420 —/07/77 FIR 151 59 64 Buckden, Wharfe 11 SD943760 =1:168 0-419 12/07/75 FJR la 24 67 Byrness, Rede 1 NT770022 1-451 0-320 22/07/62 GAS 2a Sl 67 Kielder Burn 2 NY649955 1-426 0-340 06/05/62 GAS 4a 30 70 ~—‘Irthing. Gowk Bank 4 NY682737 1-412 0-407 16/09/63 GAS 6a 36 79 Tima Water 6 NT283107 = 1:140 0-369 09/08/03 RWMCorner l6a 50 64 Ribble 12 SDITSTS9> Me355) 0:39 1027/09/03" FIR 17a 16 64 Ribble 12 SD778789 1:257 0-406 02/09/03 FJR 18a 58 64 Ribblehead SLQ 8 SDIBTS4a 2098 103638 03/09/0035) FUR: 20a 36 70 ~—=—sIrthing, below Butterburr 13. NY6873 1:196 0386 —/—/90 FJR 50a 48 70. Churnsike bridge, Irthing 23. NY6876 1517 0-406 —/—/91 FJR 26a 47 67 Whitehill quarry pond 16 NY701777 = 1-222 )3=60:353, »3—- —/08/02. T&D Hardy 29a 17 67 White Kielder Burn 19 NY677989 1-486 0-303 13/08/67 GAS 28a 4] 67 N Tyne, Charlton 18 NY809846 1:228 0-319 11/08/63 GAS 58 a 16 64 Ribble 7 SDI82786) > lca58)(0:403) 15/09/03) FIR 22a 38 64 Ribblehead SLQ 8 SD773784 1-167 0:381 03/09/03 FJR DS) 62 64 Ribblehead SLQ 8 SD772785 =: 1:287 =—-0-404_~=—- 03/09/03. FJR 33a ail 64 Ribblehead Quarry 3 SD7678 1:277° 0-405 04/09/04 FJR 34a 40 64 Ribblehead Quarry 3 SD7678 1:272 0-392 04/09/04 FJR 35a 33 64 Ribblehead Quarry 3 SD7678 1:262 0-415 04/09/04 FJR 36a 43 64 Ribblehead Quarry 3 SD7678 1:381 0-363 04/09/04 FJR Sima 66 64 Ribblehead Quarry 3 SD7678 1:160 0-383 04/09/04 FJR 46a 87 64 Ribblehead Quarry 3 SD7678 1:136 0-369 04/09/04 FJR 3a 68 64 Ribblehead Quarry 3 SD7678 1-405 0-373 04/09/04 FJR 13) -& 64 64 Ribble 7 SD782785 =1:125 0-456 15/09/03. FJR ID 2 49 64 Ribblehead SLQ 8 SD773784 1:069 0-366 03/09/03 FJR 56 ? 35 64 Ribblehead SLQ 8 SD773784 (0:884 0:365 ~==112/07/03 FIR 24? Vi 64 Ribblehead Quarry 3 SD7678 1-207 0-455 03/09/03 FJR Der, 55 70 ~—‘Irthing, Shank End 14 NY6876 1:303 0-424 —/—/91 FJR Mey th 45 70 ‘Irthing. Churnsike bridge 15 NY6873 1:142 0-424 28/09/75 FJR Dy 2S 67 Bells Braes, Irthing 17 NY691724 1:272 0-416 27/09/64 GAS Code — a = plants identified as subsp. austriaca, m = plants identified as subsp. mamillata, i = plants belonging to the intermediate group with high indices for bundle number and fruit density, ? plants which do not fall readily into any category , *s.s. stylopodium shape, *r.s.w. relative stylopodium width 320 G. A. SWAN & A. J. RICHARDS The mean and standard error of these characters were calculated for each shoot. In addition, for each shoot the following characters were recorded: e Number of vascular bundles e Stem diameter after soaking in hot water e Number of mature fruits in spikelet e Length of spikelet e Stomatal length (measured by compound microscope x400 from epidermal peels of dried material soaked in warm water, and mounted in tap water). RESULTS Across all 50 shoots there was_ highly significant variation between individuals for both stylopodium shape (s.s.: F = 17-48, P<0:0001) and relative stylopodium width G@siw:s BE = 93972, P<0:0001)) For ‘sis sthe mean reading varied between individuals by a factor of almost two, from 0-881 to 1-558, and for r.s.w. the mean reading varied from 0-303 to 0-496. There was a significant negative relationship between s.s. and r.s.w. (Fig. 1) as might be expected because these characters share the attribute of stylopodium width in which r.s.w. tends to increase and s.s. tends to decrease. The regression between these characters is identical for Yorkshire (v.c. 64) and “Borders” (vec. 67, 70) material but, although there is some overlap between the “Borders” and about one third of the Yorkshire material, the majority of the Yorkshire material has a lower average s.s. and a higher average r.s.w. than any of the “Borders” material. Taken as a whole, the variation in mean readings of both characters is continuous, showing no obvious discontinuities. We observed exceptional variability within shoots for both stylopodium shape and relative stylopodium width which, for both attributes, usually varied by a factor of two or more so that most specimens would have some fruits which were typical for subsp. austriaca and some typical for subsp. mamillata (for each of these attributes). For two individuals, both identified as subsp. austriaca, we investigated the relationship between these attributes and the position of the fruit on the spike. For individual 50 (from v.c. 70), an extreme ‘austriaca’ phenotype of mean s.s. (1:517), there was a significant relationship between fruit position and s.s. which tended to increase up the spike (F = 7-275 P—0:010) Gis 2) all the fruits on this plant fell within the range of subsp. austriaca for s.s.. However for indivi- dual 46 (from v.c. 64) with a mean s.s. of 1-136, there was no relationship between s:.s. and fruit position (F = 1-25, P = 0-267), and s.s. values of individual fruits varied from 0-64 (typical for subsp. mamillata) to more than 2-0 (an extreme value, even for subsp. austriaca). We found no relationship between fruit position on the spike and r.s.w. (F = 1-87, P= 0-178 for individual 50; F = 2:27, P = 0-135 for individual 46). Almost all specimens examined had fruits with either four or five bristles and an average bristle number in the range 4-5-5-0. Only one specimen, number 5 from Lodge Pond, Settle (Table 1), deviated from this, having an average of 5:75 bristles (n = 18 fruits, four of which had seven bristles). In this character as well as for s.s., this specimen more or less accorded with Hayek’s (1910) concept of E. mamillata. Stomatal length varied relatively little, from 44 um to 58 um, and showed no significant relationship with any other character. The number of fruits in a spikelet was expressed as fruit density, by calculating the number per cm. This figure is usually higher, above 80, for shoots with an intermediate average stylo- podium shape between 1-0 and 1:2 which is typical for neither taxon. Vascular bundle number also tended to be greatest for such intermediate individuals from v.c. 64, averaging over 20:5 (Fig. 3). DISCUSSION Based on an extensive biometric study from most of the range of FE. mamillata agg., Gregor (2003) suggested that the morphological limits of the subspecies with respect to stylopodium characters are best given as: Gregor considered that specimens with an r.s.w. Of more than 0-5 can be attributed to subsp. mamillata and those with an r.s.w. of less than 0-4 to subsp. austriaca, although many of the specimens he attributes to subsp. mamillata in his Figure 4 and the appendix to his paper (Gregor 2003) have an r.s.w. that falls ELEOCHARIS MAMILLATA 321 0.50 issp. mamillata 0.45 es ~ “n \ eae Geko Ry — eet a nee = ie — ae sen Poke = 0.40 inter mediate thister-"__ ™ . ‘ Fanon : Epa 0.30 (hes 0.9 110) il Wh ie at ee 1.6 FIGURE 1. Scatter diagram of average stylopodium shape (s.s.) for a shoot (x-axis) against average relative stylopodium width (r.s.w.) (y-axis). Symbols: circles, samples from Borders populations (v.cc. 67, 70); squares, samples from N.W. Yorks (v.c. 64); diamond, samples from Selkirk (v.c. 79). Regression lines are shown: solid, Borders samples; heavy dashing, N.W. Yorks. Ellipses encircle (from the left) samples considered to represent subsp. mamillata, all from v.c. 64; samples representing the ‘intermediate cluster’; and all the samples from the “Borders” region (v.cc. 67, 70) with one exception. ee 0 30 40 a0 0 10 FIGURE 2. Scatter diagram of position of fruit on spike (x axis) against individual fruit stylopodium shape (s.s.) for individual 50. The regression line is shown. This relationship is significant (F = 7:27, P = 0-010). G. A. SWAN & A. J. RICHARDS O,t ae 0.9 1.0 iad 1 thn) es ile 1.6 FIGURE 3. Scatter diagram of average stylopodium shape (s.s.) for a shoot (x-axis) against average fruit density on a spikelet (y-axis). Symbols: circles, samples from “Borders” populations (v.cc. 67, 70); squares, samples from N.W. Yorks (v.c. 64); diamond, samples from Selkirk (v.c. 79). Smoothing lines of best fit are shown: solid, Borders samples; heavy dashing, N.W. Yorks. Plants considered to represent the intermediate cluster are encircled. Data was missing for some plants in Fig. | and these are excluded. I , intermediate cluster Seo eee | me Ez , oe 1 0.8 0.9 1g) ei ee lst Ie; eS 1.6 FIGURE 4. Scatter diagram of average stylopodium shape (s.s.) for a shoot (x-axis) against number of vascular bundles on that shoot (y-axis). Symbols: circles, samples from Borders populations (v.cc. 67, 70); squares, samples from N.W. Yorks (v.c. 64); diamond, samples from Selkirk (v.c. 79). Smoothing lines of best fit are shown: solid, Borders samples; heavy dashing, N.W: Yorks. Plants considered to represent the intermediate cluster are encircled. Data was missing for some plants in Fig. 1 and these are excluded. ELEOCHARIS MAMILLATA 328 between 0-4 and 0:5. In fact, no British plant measured had an average r.s.w of greater than 0-50. However, eight individuals from York- shire had an average s.s. of less than 0:97 which Gregor considered typical for subsp. mamillata. These plants also gave a relatively high r.s.w. reading of more than 0-429 (Fig. 1). A further plant from Selkirk also falls within this group. All the plants from the Borders region (vcc. 67 and 70) had an s.s. of more than 1-10 and so fall within Gregor’s definition of subsp. austriaca for this character. For all (except five) the r.s.w. falls below 0-40 which is also diagnostic of subsp. austriaca according to Gregor. It is clear that it is necessary to measure a large number of fruits from all parts of the spikelet before a consistent average reading is obtained. Measurements of single fruits, or even much larger samples from the same part of the spikelet might give highly misleading readings. For instance, the average s.s. for the bottom half of spike 50 was 1-39. and that for the top half was 1-64 (n = 24 in each case). Gregor (2003) measured either three or five fruits per spike and does not state from which part of the spike these were taken from. It is our experience that such small samples would be liable to considerably bias average readings for s.s. and r.s.w., particularly if the part of the spike from which they were taken was not standardised. Nevertheless, when large samples are taken from the total spike, as in the present study, there is considerable Statistically significant variation between individuals. However, like Gregor, we conclude that there is no clear discontinuity in the distribution of the data across all populations which could have provided a natural basis to the delimitation of the subspecies. Also, like Gregor, we note that a proportion of individuals are intermediate in character and cannot be readily referred to either taxon. In fact, amongst our material, there is a distinct cluster of 12 shoots originating from Yorkshire with an average s.s. between 1-00 and 1:18 and an average r.s.w. between 0-40 and 0:45 which form a distinctive morphological group. This group of individuals has on average more vascular bundles than any other with a single exception (between 20:5 and 22:0) (Fig. 4), and a higher fruit density than any other (more than 80/cm) with three exceptions (Fig. 3). The only shoot from elsewhere which resembles this intermediate group comes from the Tima Water, Selkirk, but this has a much lower r.s.w. average (0:37). We are uncertain as to the significance of the high readings for fruit density and vascular bundle number for this intermediate group. Both characters have been regarded as having taxonomic significance in earlier work. Beauverd (1921) considered that a fruit density of 60-70 was typical of E. benedicta Beauverd which Walters (1953) regarded as synonymous with E. (mamillata subsp.) austriaca. This agrees well with our findings for plants classified (according to mean s.s.) as both subsp. mamillata and subsp. austriaca, but is lower than we find for our ‘intermediates’. Hayek (1910) stated that subsp. mamillata tends to have fewer vascular bundles than subsp. austriaca (i.e less than 12). Gregor (2003) found that 61% of individuals he classified as subsp. mamillata had 13 bundles or fewer. However, all our shoots except two had more than 13 bundles, and there is no clear distinction in bundle number between our material which, according to mean s.s., was classified as ‘typical’ subsp. mamillata and ‘typical’ subsp. austriaca. Indeed, the number of bundles in the eight individuals that fall clearly within the s.s. character range for subsp. mamillata for which bundle number is known, shows the complete range present, from 12:5 to 22 (Fig. 3). Strandhede (1966) considered that bundle number was proportionate to stem diameter, and thus had no taxonomic signifi- cance, although in our opinion high bundle number could be a measure of vigour. We believe two interpretations of this data are possible. Either, the intermediate individuals with high bundle and fruit density readings should be regarded as a distinct taxon, or these attributes can be considered as measures of vigour, in which case individuals with an intermediate stylopodium shape might be considered to show hybrid vigour. If we regard these intermediate individuals as a distinct taxon, they have several features in common with the little-known E. benedicta Beauverd (1921), described from Lac Benit, Haute-Savoie, at 1500 m., notably: the stylopodium shape, said to average 1-07, the relative stylopodium width, said to average 0:39, the fruit density of 80/cm and the number of perianth bristles (5). However, E. benedicta, regarded by Walters (1953) as a synonym of 324 G. A. SWAN & A. J. RICHARDS austriaca, was stated to have a low vascular bundle number (averaging 12, rather than more than 20 in our ‘intermediates’). As the high bundle number is one of the main attributes of the latter, we hesitate to identify it with E. benedicta. With regard to the vigour hypothesis, this assumes that two previously isolated demes have come together in Yorkshire and have hybridised. Some support for this hypothesis can be derived from the finding that three of the four Yorkshire sites from which three or more individuals were collected (Ribble 7, Ribble 12, and Ribblehead SLQ 8) contained spikes showing stylopodium characters of subsp. austriaca, subsp. mamillata and the intermediate. We have _ considered the possibility that plants of intermediate phenotype and characteristics of vigour are allotetraploids, but the only chromosome counts recorded are diploid (2n = 16, 18) (Walters 1953, Strandhede 1966). Stomatal length commonly differs between related diploids and tetraploids (e.g. Stebbins 1950) and there is no suggestion from stomatal length that intermediates are tetraploid. Eleocharis fruits are readily dispersed, especially by ducks (Gregor & Barth 1998). Some of the localities for E. mamillata agg. in Britain are artificial (i.e. quarries, gravel pits and reservoirs) and most are impermanent. Whether or not this group has persisted within these islands for much of the post-glacial period, it is quite likely that immigrant wildfowl have mediated repeated colonisations of both subspecies over thousands of years, as is suggested by the Yorkshire sites. In this context, it is perhaps remarkable that most individuals in the Borders populations are clearly referable to subsp. austriaca, suggesting that they may have originated from relatively few founder events. The question remains as to the correct name for the plants from N.W. Yorks and Selkirk which have a stylopodium shape (s.s.) within the range given by Gregor (2003) for subsp. mamillata. We suggest that it would be unacceptable to record this taxon from the British Isles for the first time on the basis of this single character. Using relative stylopodium width (r.s.w.), none of these plants fall within Gregor’s stated limit for subsp. mamillata (although this is also true of many of his so-called subsp. mamillata), and they have a vascular bundle number well in excess of that considered to be typical for this taxon. Except for one specimen, all of the material which we examined appeared to have an average perianth bristle number of five or less (i.e. typical of subsp. austriaca) including individuals such as number 31 which have an average s.s. typical of subsp. mamillata. Only individual 5 had an average bristle number typical of Hayek’s concept of subsp. mamillata. Gregor (2003) shows that individuals assigned to subsp. mamillata can have the full range of average bristle number recorded, from 4:0 to 7:0. Although he found that no individual of subsp. austriaca had an average of more than 5-5 bristles, this character seems at best to be of very limited value. Unfortunately, very few characters seem to separate these two taxa reliably and those that do so (s.s. and r.s.w.) are both based on the same basic character: the shape of the stylopodium. Nevertheless, our material is not typical of subsp. mamillata for r.s.w., perianth bristle number or vascular bundle number, and further work is needed before it can be safely assigned to ssp. mamillata. It has been suggested that the ecology of the two subspecies differs considerably, at least within Scandinavia. However, some of the British sites are man-made and appear to differ considerably from the natural localities in which they typically occur abroad, although Gregor (in litt. to F. J. Roberts) notes that in Germany both taxa occur predominantly in derelict ponds. As yet, there are no ecological data from British sites that can be compared with that gathered by Strandhede (1966) and Gregor & Barth (1998). We conclude that plants in which the clear majority of fruits have a stylopodium longer than wide should be recorded as E. mamillata subsp. austriaca. All other British material should at present be referred to as E. mamillata agg: ACKNOWLEDGMENTS We would like to thank F. J. Roberts (FJR) for his very considerable contribution both in collecting material and in later discussions. Thanks are also due to the late Mrs O. M. Stewart, Dr R. W. M. Corner, and Mr T. and Mrs D. Hardy for collecting material, and to Dr M. Swan who was involved in the location and collection of much of the material. ELEOCHARIS MAMILLATA 325 REFERENCES BEAUVERD, G. (1921). A propos d’une nouvelle espéce européenne du genre Eleocharis. Bulletin de la Société Botanique de Genéve ser. II (2), xiii, 245-265. CORNER, R. W. M. (1975). Eleocharis austriaca Hayek new to Scotland. Watsonia 10: 411-412. GREGOR, T. (2003). Eleocharis mamillata — distribution and infraspecific differentiation. Folia Geobotanica 38: 49-64. GREGOR T. & BARTH, U. (1998). Die Weichstengelige Sumpfbinse Eleocharis mamillata in Hessen. Natur und Museum 128: 113-124. HALLIDAY, G. (1997). A Flora of Cumbria. Centre for North-West Regional Studies, University of Lancaster. HAYEK, V. A. (1910). Schedae ad floram stiriacum exisccatum. 19 und 20. Lieferung, Wien. HEGI, G. (1966). I/lustrierte Flora von Mitteleuropa, 3rd ed., 2 (1): 61-64. Carl Hanser Verlag, Munchen. Hess, H. E., LANDOLT, E. & HIRZEL, R. (1967). Flora der Schweiz und angrenzender Gebiete 1: 397. Birkhauser, Basel and Stuttgart. ROBERTS, F. J. (1977). Eleocharis austriaca in N. Yorkshire. B.S.B.I. News 15: 16-17. STEBBINS, G. L. (1950). Variation and Evolution in Plants. Columbia, New York. STRANDHEDE, S.-O. (1966). Morphological variation and taxonomy in European Eleocharis, subser. Palustres. Opera Botanica 10: 1-187. STRANDHEDE, S.-O. & DAHLGREN, R. (1968). Drawings of Scandinavian plants 9-10. Eleocharis R. Br. Botaniska Notiser 121: 1-10. SWAN, G. A. (1993). Flora of Northumberland. Natural History Society of Northumbria, Newcastle upon Tyne. WALTERS, S. M. (1953). Eleocharis mamillata Lindb. fil. and allied species. Bericht der Schweizeriscen Botanischen Gesellschaft 63: 271-286. WALTERS, S. M. (1963). Eleocharis austriaca Hayek, a species new to the British Isles. Watsonia 5: 329-335. WALTERS, S. M. (1980). Eleocharis R. Br., in T. G. TUTIN, V. H. HEyYwoop, N. A. BURGES, & D. H. VALENTINE eds. Flora Europaea 5: 281—284. Cambridge University Press, Cambridge. (Accepted May 2006) | , fh i } | | } if Watsonia 26: 327—338 (2007) 327 The University of Hull Herbarium R. MIDDLETON*% and J. R. MIDDLETON Department of Geography, University of Hull, HU6 7RX ABSTRACT The University of Hull Herbarium was transferred from the Botanic Gardens to the Geography Depart- ment in 1993 by which time almost all documentation, including the catalogues, had been lost. Over the following five years a computerised catalogue was created by transcribing the labels of over 18000 British vascular plants. The larger part of the collection was the teaching and research collection (HLU) created by Prof. Ronald Good in the period between his appointment in 1928 and retirement in 1959. An additional collection of 1262 specimens had been acquired by the University in 1983; this is now believed to be the Herbarium of the Hull Technical College (HLL), assembled by Thomas. Stainforth. Although it was known that Good had acquired the herbarium of Dr John Fraser (1820-1909) of Wolverhampton, analysis of the new catalogue has revealed the presence of other substantial collections. Over 110 specimens from the herbarium of Rev. W. O. Newnham (1825-1893) have been located and almost 700 19th century specimens from the herbarium of the Barrow Naturalists’ Field Club, recorded as having been destroyed in the 1930s. HLU also contains 676 plants from the undocumented herbarium created by Rev H. W. G. Kenrick (c.1863—1943). Geographical coverage of the material is wide but vice-counties 9, 37, 39, 61 and 69 are particularly well represented. The catalogue has now been made public via the internet and positive feedback has enabled the correction of many transcription errors. INTRODUCTION The University of Hull Herbarium was established by Prof. Ronald Good shortly after his appointment as head of the Botany Department at the newly founded University College in 1928. The collection of British material, which seems to have been primarily intended as a teaching resource, was added to until Good’s retirement in 1959 and then only sporadically into the 1960s. In 1993 the herbarium was moved from the University’s Botanic Gardens in Cottingham, where it had been since about 1975, to its present home within the Department of Geography. In the Geography Department it is now housed along with an extensive collection of South-east Asian material collected by Dr John Flenley *e-mail :r.middleton @hull.ac.uk and others. Unfortunately most of the original catalogues and any documentation that may have once been associated with the material had been lost. This paper describes the re- creation of the catalogue of British and Irish vascular plants and establishes the origins of Good’s original University Herbarium. It has not yet been possible to assess the many thousands of still largely un-mounted European specimens, that originally formed part of the Fraser collection, or the mosses, lichens and fungi. In 1997 a start was made to build a new computerised catalogue of the British material. After examining the alternatives, it was decided that the most convenient format for the catalogue would be in the form of a Microsoft Access relational database. This provided a widely used and accepted standard with the option of exporting the data in many other formats, as required. Rather than using direct data entry, a Borland Delphi application was written to ensure that the nomenclature conformed to recognised standards (Kent 1992; Clement & Foster 1994; Ryves et al. 1996). This was particularly important as, although the collections were organised according to Dandy (1958), many taxa and hybrids did not fit well into his system. It also ensured the correct spelling of the taxon name field in the database. Several small grants enabled all of the British vascular plant material to be catalogued and some of the previously unmounted material to be mounted. Much of the data presented here has been derived from analysis of this computerised index. The value of the complete catalogue has proved to be greater than the simple sum of its individual parts and by sorting the records by collector, collection date and locality it has been possible to extract information that was not apparent from the original taxonomic ordering. Many virtually illegible labels have been deciphered by comparison with others. At the present time there are over 18000 catalogued vascular plants in the collection. Of these, approximately 14000 plants can be assigned a collector, 16500 a locality and 17000 a collection date. 328 R. MIDDLETON & J. R. MIDDLETON DESCRIPTION OF THE BRITISH MATERIAL The University Herbarium was established by Prof. R. Good soon after his appointment as Lecturer in Botany in 1928 and it is first mentioned in the Hull University College Calendar for 1931-1932. The most recent pub- lished synopsis of the material has been the report resulting from the Northern Universities’ Collection Survey (Arnold-Foster 1993). The entry for the Hull herbarium is brief and mistakenly states that all of the material was collected between 1884 and 1971. In fact more than a third of the material is earlier than this. The draft report of a survey of a visit for this survey of collections, sent to the University in 1991, still survives and it is clear that the misapprehension went unnoticed and un- corrected. This draft report does indicate that at that time there were five handwritten catalogue volumes for the British plants and a further one for the European material. These appear to have been lost around the time of the transfer to the Geography Department. The British vascular plants had _ been organised into two sections, the larger part consisting of over 16000 mounted specimens grouped into folders by species, the folders then being grouped into genera covers. The folders were ordered according to Dandy (1958) and stored in wooden cupboards. With this were several boxes of unmounted British material, largely willows, interleaved with late 19th century newspaper. The second section was simply labelled the “Wilson Collection”, consisting of 1263 mounted specimens arrang- ed into folders by genus and then grouped into conveniently sized bundles. The larger collection is the core of Good’s Herbarium and its primary purpose seems to have been a taxonomic resource for teaching and research within the Botany Department. As other collections were acquired, they were merged with this and consequently lost their integrity. The description of HLU given by Kent and Allen (1984) “Formed by past and present members of staff and students” 1s basically correct but it hides the presence of older material that was acquired by donations and possibly purchase. A card index to the East Yorkshire material has survived. This was created at some time in the later 1950s by Miss A. B. Mowatt (F. E. Crackles, pers. comm.), who worked in the herbarium for a while. It seems to represent preparations for an unpublished Hand-list of the Flora of the East Riding of Yorkshire prepared by Good and Crackles at this time (Crackles 1990). Surprisingly, this index provides evidence for the origin of the “Wilson Collection”. Each taxon card carries a list of early v.c.61 records, presumably taken from the literature, followed by two lists of localities, one labelled “HUH” and the other “Tech”. After the former is a list of records taken from the University Herbarium and the latter corresponds to plants in the “Wilson Collection”. “Tech” seems to refer to the Hull Technical College, the precursor of the Hull College of Higher Education. Kent and Allen (1984) record that A. K. Wilson’s herbarium was held in the Hull Technical College Herbarium, HLL. The only reference to its acquisition is to be found in the draft report made in preparation for the Northern Uni- versities’ Collection Audit (Arnold-Foster 1991). This report records that E. K. Wilson’s (sic) British and European Flora was acquired as a gift in 1983, although it is not recorded from where. It seems safe to conclude that the present University herbarium now incorporates both HLU and HLL. The HLL material covers a much more restricted period than that in HLU. If the 80 sheets of Hull plants submitted for the 1864 British Botanical Competition, which clearly form a discrete collection, are excluded, only 60 specimens pre-date 1930 and none is later than May 1939. Most of the early specimens are attributable to exchange club material (Arsene, Melville, etc) or local amateurs (Boult, Robinson). This matches well the period that Thomas Stainforth was lecturer in Botany and Zoology at the Technical College and he may have established and developed the collection. Stainforth had strong links with the Hull Scientific and Field Naturalists’ Club (HSFNC) which he served, at various times, as secretary, treasurer, editor and chairman (Walsh 1944). There is little duplication of specimens in HLL and it seems to represent a deliberate attempt to build a_ teaching collection. CONTRIBUTORS Over 300 collectors have contributed to the current herbarium at Hull University, those with the greatest number of specimens are listed in Table 1. Ronald Good has by far the most specimens with almost 20% of the collection being of his own material. A THE UNIVERSITY OF HULL HERBARIUM 329 TABLE 1.THE MAJOR CONTRIBUTORS TO THE HERBARIUM AT HULL UNIVERSITY HLU HLL Good, R. D’O. 3576 - Wilson, A. K. 237 784 Fraser, J. 1258+ - Kenrick, H. W. G. 676 - Wilkins, D. A. 297 - Lewis, R. DT - Baynes, J. & T.C. 343 - Fisher, J.* DS) - Hardy, A. C. 723) - Rob, C. M. 70 156 Ley, A. 173 - Gardiner, W. 173 - Worsdale, A. F. 140 - Shoolbred, W. A. 27 - Bennett, A.* 120 - Baker, J. G.* mS - Newnham, W. O. 110+ - Brotherston, A. 95 - Baxter, T. 62 = Smith, R. L. 53 2 * Indicates approximate numbers resulting from similarly named collectors HLU = the original Hull University Herbarium HLL = the former Technical College Herbarium personal collection of over 800 specimens, collected prior to his appointment, may well have formed an early core. An interesting and unsuspected addition in the early years is a set of over 200 plants collected in the Harrogate area in 1914 and attributed to A. C. Hardy. These specimens were almost certainly coll- ected by Sir Alister Hardy, who was appointed as head of the Zoology Department in 1948. Although Sir Alister Hardy later attained acclaim for his work on marine plankton, he had early botanical interests, entering Oxford University in 1914 to study forestry ento- mology. Other collections were acquired, the most notable of which is the herbarium of John Fraser of Wolverhampton. Over 1000 specimens in the British collection can be assigned, with confidence, to Fraser. This includes a series of around 300 Staffordshire plants, collected in 1864 for the British Botanical Competition organised by the South Kensington Horticultural Society. Edees (1972) notes that Fraser won a silver medal with a collection of 530 species; clearly some of these must be elsewhere or lost. There are other notable collections that once formed part of Fraser’s herbarium. More than 110 specimens have been identified in HLU which may be assigned, with certainty, to Rev. William Orde Newnham (1825-1893). There is no doubt that these specimens at one time formed part of John Fraser’s herbarium as there are examples where Fraser has altered his own printed labels to read “Ex Herbario Newnham”’. Most specimens, however, are identifiable only by the initials WON or even just N and there is even a note, in Fraser’s hand, stating “Newnham specimens unlabelled”, suggesting that some of the unattributed and unlocated specimens in his herbarium may also have originated with this collector. Although Desmond (1994) records the existence of material collected by Newnham in CGE (Cambridge University) and MANCH (Man- chester Museum), none of his British material is catalogued in the former and only a few specimens (previously attributed to Newsham) are to be found in the latter. It would appear that HLU may hold the bulk of Newnham’s. surviving British material. All dated Newnham specimens cover the period 1843-1846, when he would have been a Cambridge undergraduate, and most are from the vicinity of Chesham, Bucks. There are at least another 20 Buckinghamshire records from 330 this period that will almost certainly have been collected by Newnham. Although the records show that both William Orde Newnham and John Fraser were members of the Botanical Society (Desmond 1994), there is no other known link between the two men which might explain the transfer of large numbers of plants. However, examination of census records reveals that by 1861 W. O. Newnham’s brother, Christopher Atkinson Newnham, was a doctor in Wolverhampton. As a fellow medical practitioner, C. A. Newnham would certainly have been well known to John Fraser as both men resided in the town for the following thirty years. It seems likely that W. O. Newnham’s specimens arrived in Fraser’s herbarium via his brother. The 62 plants of T Baxter of Worcester may also have once been part of Fraser’s collection. Fraser’s extensive collection of European plants is also housed within the Herbarium but remains un-catalogued (much of it still being un-mounted). A significant component of the HLU material is that derived from the Botanical Exchange Clubs. The HLU section has around 760 such specimens ranging in date from Baker’s 1860 specimen to Druce’s 1931 contribution. This is significant as although Fraser was a member of the Botanical Exchange Club and such specimens would be expected, he died in 1909 and _ seventy specimens post-date this. There must be specimens from another herbarium that were incorporated after 1930. There is significant material from the Furness district of Cumbria. This was largely collected by Mr T. C. & Mrs J. Baynes and Mrs J. Fisher during the period 1876-1880. Examination of the Transactions of the Barrow Naturalists’ Field Club for this period reveals that these collectors formed a nucleus of an active botanical section. Many of _ the specimens now in HLU were collected on the Club’s organised field excursions. In 1878 Mrs. Joseph Fisher is recorded as obtaining a prize for “... the best herbarium of flowering plants, LEPLEesenting. Not less Methane 0 species...” (Anon 1878). How this material came to be in Hull was unclear until a note was found on an_ unattributed specimen’ of Cephalanthera rubra stating that it “... came with the Ferguson plants from Barrow ...”. Dr J M Ferguson was an Assistant Lecturer in the Hull Botany Department from 1930 to 1932 and of the four specimens she left in the R. MIDDLETON & J. R. MIDDLETON herbarium one is from Walney, collected in 1931. Further examination of the Barrow Naturalists’ Field Club Proceedings has revealed that the Club’s collections were moved from their premises at Cambridge Hall around 1932 where they had suffered seriously from damp and dirt. The botanical collection had fared particularly badly and was “...In such a condition, that the committee decided to destroy it, but on the advice of Dr Judith Philip, M.Sc. Sheffield, daughter of Mr. T. J. Ferguson, the whole collection was sent just as it stood to the Hull University College. Dr. Ronald D’O Good, M.A., Cambridge, Professor of Botany at the College, in acknowledgement, said that although the specimens had_ perished, the labels still remaining, were of great value ...” (Christian & Richardson 1933) It seems that the material was not in such a poor state as suggested by this, and that a significant number of specimens were re- mounted and added to the embryonic HLU. This misconception is propagated further when later proceedings report again that the specimens had “mouldered _ into dust” (Christian 1935) but that the record cards had been sent to Hull University. Although there is no specific record of the size of the Barrow herbarium, it is possible to estimate it at around 800 specimens. The Barrow Club’s Botanical Report for 1877-— 1878 states that Mrs Baynes had donated 356 specimens and that over 200 had _ been contributed by other members. HLU contains about 340 specimens attributed to J. or T. C. Baynes, for this period, and about 240 for Mrs J. Fisher. A later report (Anon 1902) notes that Mr John Hosking, a long-standing member, had donated over 200 specimens to the Club’s herbarium in 1896. HLU only holds 59 specimens that can be attributed to J. Hosking but it is of interest that of the 49 plants specifically mentioned in the report, 43 can be positively identified in HLU. The best estimate that can be made of the surviving Barrow material is around 680 re-mounted specimens. This would suggest that, despite pessimistic reports, over 80% of the material may have survived. The 676 specimens collected by H. W. G. Kenrick between 1889 and 1939 are of unknown origin. Little is known of this collector as a botanist but it is possible to THE UNIVERSITY OF HULL HERBARIUM 3311 deduce from the labels that he was Rev. Henry William Gordon Kenrick (c.1863—1943), an Anglican Priest, who spent much of his min- istry in London. Confirmation of this is prov- ided by a 1903 Dandelion specimen from St John’s vicarage, Hammersmith. He was born in Bangalore, India and completed an MA at Cambridge (Venn 1954). Coincidentally, be- tween 1888 and 1890 he was attached to Holy Trinity Church, Hull, and his earliest specimen was collected at Bridlington, East Yorkshire during this time. He appears to have main- tained a botanical interest through most of his life and his final specimens were collected in Dawlish in September 1939. It is noteworthy that there are no grasses in the collection and only a single rush. It seems likely that part of the collection was lost or is elsewhere. As is to be expected in a provincial coll- ection, many of the collectors do not figure in standard botanical directories (Britten & Boulger 1931; Desmond 1994). Both sections of the herbarium contain significant amounts of material collected by local amateurs. During the 1930s, when both the University and Technical College collections were being actively built, local and Yorkshire naturalists including A. K. Wilson, C. M. Rob and A. E. Wray contributed to both collections. The minute books of the HSFNC for this period show that Stainforth was the president of the Club in 1929, the year that Good applied for membership, and that A. K. Wilson became their Botanical recorder in 1931. During this period Ronald Good gave regular lectures to HSFNC and their meetings were often held in the University. The major exceptions to this pattern are the 168 specimens collected by G. A. Glister and the 300 or so specimens coll- ected by Good himself in this decade, which are found only in HLU. Although the Diatomaceae recorder for HSFNC, Glister obviously had close association with the University College as the Botany Department is given as his address in Club records. Even Stainforth, more an entomologist by inclin- ation, has 8 specimens in HLU. Considering that only 25 of Stainforth’s plants are in HLL, the flow of material seems to have been a little one sided. The almost equal distribution of Royston L. Smith’s Cardiff alien specimens between the two herbaria is undoubtedly a result of donations from A. K. Wilson. Wilson was extremely interested in dock aliens (Wilson 1938) and it is known that the two collectors exchanged material. After 1940 Good’s connection with the HSENC seems to have been a little more distant, although he was eventually made an Honorary Life Member of the Club. Approaching 70% of the Wilson specimens in HLU date from 1939 or later. It is not clear whether there was a constant trickle of don- ations or whether the post-war part of Wilson’s collection was donated to the University at some time. The latter may be more likely as an undated (1950s) letter from Mr Foster, HSFNC’s secretary at the time, in response to a request from Good concerning the whereabouts of Charles Waterfall’s herbarium, suggests that Wilson might know (DX/14/4). The tone of the letter makes it clear that Wilson was not now an active member of the club and that Foster was unaware that Good and Wilson may be acquainted. Robert Lewis provided a link between Good and Wilson in the early 1950s. Lewis was an active member of the HSFNC and donated almost 300 specimens to HLU between 1949 and 1955, after which he moved from the area. There are 18 specimens in HLU attributed jointly to Wilson and Lewis, the majority resulting from a visit to Westmorland in June of 1954. Lewis also botanised with D. A. Wilkins, Good’s assistant at the time. Wilkins’ 297 specimens were all collected during 1952 and all, bar one, are from v.c.6l. Angus Worsdale, another HSFNC member also made a significant contribution of local specimens during this period. Dr Eva Crackles, HSFNC member, student of Good and graduate of Hull University, is also well represented in HLU. Almost fifty specimens, the majority of which are local, were contributed during the 1950s. A further 40 specimens, mainly from 1981, which formed a separate collection, have recently been integrated with the main body of speci- mens. Crackles made extensive use of the university herbarium in the preparation of her Flora of the East Riding of Yorkshire (1990). Unsurprisingly, there are many specimens contributed by students and staff of the University. Usually there are only a few per collector but an exception to this is the material collected by the student Andrew Caldicott, mainly in 1967. These 73 specimens, mainly from Essex, form the last substantial addition to the herbarium. 332 R. MIDDLETON & J. R. MIDDLETON 200 Kilometres FIGURE 1. Geographical distribution of specimens in the herbarium (HLU and HLL) from within Great Britain. Each dot represents one specimen but is located randomly within the vice-county. - THE UNIVERSITY OF HULL HERBARIUM 333 GEOGRAPHICAL DISTRIBUTION OF SPECIMENS Figure | illustrates the spatial distribution of the specimens which can be assigned to the vice-counties of Great Britain. It is clear that there is a very uneven spatial distribution, a function of the dominance of certain collectors. As to be expected, South-east York (v.c. 61) is extremely well represented with almost a third of all specimens originating within the vice-county. Over 2500 of these specimens were collected by Prof. Good with 87% being added in the 1950s. A. K. Wilson made the next largest contribution, almost 1800 speci- mens, all but 30% of which were collected before 1950. Staffordshire (v.c. 39) is the next best- represented county. Although only 258 of these specimens are undoubtedly John Fraser’s, the great bulk of the 1059 bearing no collector’s name are almost certainly from his herbarium. The actual number of Staffordshire specimens will be greater than indicated as many of Fraser’s labels are illegible. The adjacent vice- counties are also reasonably represented, particularly Worcester (v.c. 37). Almost 100 of the 760 specimens are known to have been collected by, or are from the herbarium of John Fraser. Although these plants are supplemented by 36 specimens collected by T. Baxter of Worcester, the great majority of those without a known collector are probably Fraser’s. Numerically the collections of plants from Westmorland (v.c. 69) are worthy of mention. They are dominated by the Baynes, Fisher and Hosking material of the 1870s described earlier, but there are significant contributions from H. W. G. Kenrick, who appears to have made several summer visits to the area. The Dorset specimens (v.c. 9) are also of signifi- cance. Around 80% of the Dorset material was collected by Prof Good, the bulk from the period 1914-1939. These will be part of the collection made for his Dorset Flora (Good 1948). Some manuscript maps and draft material for this work are among the little surviving herbarium documentation. The 260 specimens for Mid Perth (v.c. 88) are un- surprising and they comprise the contributions of over 25 individuals, most of whom were visiting Ben Lawers. TAXONOMIC COVERAGE As is to be expected of a university depart- mental collection, coupled with Ronald Good’s own interests, which were often closer to biogeography than to taxonomic botany, HLU covers the British flora reasonably comprehensively with representatives of 2001 taxa. The extensive incorporation of earlier herbaria into HLU has, however, resulted in better representation in groups which were of particular interest to their original owners. Until 2002 Fraser’s Salix collection remained un-mounted and still interleaved with 19th century newspaper sheets. Of the 524 Willows in HLU, probably a third result from Fraser’s own collectings. Andrew Brotherston is also strongly represented and an interleaved letter shows that Fraser had requested specimens from him. Other notes show that Fraser was also corresponding with W. H. Beeby and E. F. Linton about Willows. Both Rubus and Rosa are well represented in HLU, with 269 and 264 specimens respectively. Much of this material appears to have originated from Fraser’s herbarium where Bloxam, Bagnall and other 19th century collectors feature strongly. HLL, despite containing less than a tenth of the specimens, contains representatives of almost half as many taxa as HLU. This is consistent with the hypothesis that it was established as a teaching rather than research collection. Alien species are particularly well represented in HLL with 55. specimens collected by R. L. Smith, mainly from Cardiff, supplementing Wilson’s material from the Hull docks. Table 2 lists the specimens from HLU and HLL which appear on the Red Data List as being either extinct in the wild or critically endangered. In addition HLU holds specimens of 56 taxa classified as endangered and 16 are to be found in HLL. DISSEMINATION OF INFORMATION It was the intention, from the start of this project, to make the catalogue freely available to as wide an audience as possible. A pilot internet service was established in early 2000 using a provisional version of the catalogue. As no commercial software suitable for the task was available at the time, a dedicated program was written in Borland Delphi. This program, which could be called from a standard web page, permitted interrogation of the catalogue by taxon, collector, locality or v.c. number, together with several options for the sorting of results. The results were returned as a web page, requiring no special software on the user’s part. The software was upgraded several times over the following two years and now 334 R. MIDDLETON & J. R. MIDDLETON TABLE 2. SPECIMENS OF TAXA APPEARING ON THE RED DATA LIST AS EITHER EXTINCT IN THE WILD OR CRITICALLY ENDANGERED HLU taxa now extinct in the wild Euphorbia peplis Euphorbia peplis Euphorbia peplis Euphorbia peplis Euphorbia peplis Euphorbia peplis Galeopsis segetum Arnoseris minima Arnoseris minima Arnoseris minima Crepis foetida Crepis foetida Filago gallica Filago gallica Filago gallica Filago gallica Otanthus maritimus Otanthus maritimus Otanthus maritimus Otanthus maritimus Otanthus maritimus Tephroseris palustris Spiranthes aestivalis Spiranthes aestivalis Spiranthes aestivalis Spiranthes aestivalis HLU critically endangered taxa Dryopteris cristata Dryopteris cristata Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Ranunculus arvensis Chenopodium urbicum Chenopodium urbicum Chenopodium urbicum Chenopodium urbicum Chenopodium urbicum Chenopodium urbicum V.C. IS) 11S 113 113 H12 i) 113 Locality Slapton Sands Dartmouth Slapton Sands Beesands Torcross Beesands Bangor Verwood Wrecclesham Crosswood, Allerthorpe Kingsdown, Deal Dartford to Greenhithe Berechurch Berechurch Colchester Bayford St Ouen’s Bay, Jersey St Ouen’s Bay, Jersey St Ouen’s Bay, Jersey St Ouen’s Bay, Jersey Wexford East Norfolk Lyndhurst Lyndhurst St Ouen’s Pond, Jersey St Ouen’s Pond, Jersey Kingston Pool Wybunbury Bog Ruddipole, Weymouth Thames Ditton Berkhamstead Maidenhead Lower Penn Waudby Cottingham Kilnsea Bewholme Burstwick Fitling Easingwold Kew Bridge Chobham Waddon, nr. Croydon Epping Forest Epping Forest Cottingham Collector Curnow, W. Ralfs, J. Ralfs, J. Waterfall, W. B. Waterfall, W. B. Marshall, E. S. Ley, A. Good, R. D’O. Nelson, G. A. Bennett, A. Linton, W. R. Tempere, J. Dufrey, J. ? Linton, W. R. Groves, H. Blow, T. B. Mathews, W. Lomax, A. E. Lind, J. Hurst ? Hanbury, F. J. Teesdale, M. J. Goss, H. Fraser, J. Fraser, J. Fraser, J. Good, R. D’O. Preston, A. W. Good, R. D’O. Good, R. D’O. [Fraser, J.] Wilson, A. K. Good, R. D’O. Good, R. D’O. Allison, J. H. Worsdale, A. F. Good, R. D’O. Good, R. D’O. Bennett, A. Crespigny, E. de Crespigny, E. de Good, R. D’O. Corrigiola litoralis Corrigiola litoralis Corrigiola litoralis Corrigiola litoralis Corrigiola litoralis Corrigiola litoralis Sorbus domestica Sorbus domestica Sorbus domestica Sorbus domestica Sorbus domestica Sorbus domestica Sorbus domestica Cotoneaster cambricus Cotoneaster cambricus Cotoneaster cambricus Cotoneaster cambricus Cotoneaster cambricus Cotoneaster cambricus Cotoneaster cambricus Eryngium campestre Eryngium campestre Eryngium campestre Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Scandix pecten-veneris Bupleurum rotundifolium Bupleurum rotundifolium Bupleurum rotundifolium Cynoglossum germanicum Cynoglossum germanicum Cynoglossum germanicum Cynoglossum germanicum Galeopsis angustifolia Galeopsis angustifolia Galeopsis angustifolia Galeopsis angustifolia Galeopsis angustifolia Galeopsis angustifolia THE UNIVERSITY OF HULL HERBARIUM TABLE 2 CONTINUED Locality Looe Pool, Helston Looe Pool, Helston Looe Pool, Helston Helston Slapton Walney Kew (cult.) Kew (cult.) Wyre Forest, Bewdley Wyre Forest Arley Castle Arley Castle grounds Arley Castle Great Orme’s Head Llandudno Great Orme’s Head Great Orme’s Head Ormeshead Great Orme’s Head Great Orme’s Head Devil’s Point Devil’s Point Devil’s Point Sydling St. Nicholas Holyport, Maidenhead Cowleigh Park ? Trysull King George Dock, Hull Goxhill Pocklington Welwick Kilnsea Warren Skidby Fitling Roosebeck Bredicot Cracombe Hill Harbury Norbury Park Norbury Norbury Tewkesbury Dorking Wauldby Millington Grange Huggate Dikes South Dalton Brantingham 335 Collector Year Beeby, W. H. 1873 Curnow, W. 1875 1876 Fraser, J. 1878 Fooks, K. 1949 Baynes, J. 1877 Baker, J. G. 1874 Nicholson, G. 1878 Jordan 1849 1853 1865 Fraser, J. 1878 1887 G. Seeb.. truncated - 1853 Baxter, T. 1855 Woodward, J. 1856 1863 Griffith, J. E. 1884 Griffith, J. E. 1889 Goulding, F. H. 1846 Hayward, W. R. 1873 Briggs, T. R. A. 1876 Good, R. D’O. 1914 Good, R. D’O. 1923 Fraser, J. 1877 Fraser, J. 1864 Wilson, A. K. 1937 Glister, G. A. 1938 Nelson, G. A. 1939 Sledge, W. A. 1946 Wilson, A. K. 1952 Good, R. D’O. 1952 Good, R. D’O. 1954 Fisher, J. 1877 Baxter, T. 1860 Fraser, J. 1877 Groves, H. 1875 Goss, H. 1886 Goss, H. 1888 1854 Morgan, J. H. 1896 Good, R. D’O. NO Sik Lewis, R. 1952 Good, R. D’O. 1953 Good, R. D’O. 1956 Good, R. D’O. 1956 336 R. MIDDLETON & J. R. MIDDLETON TABLE 2 CONTINUED v.c. Locality Collector Year Galeopsis angustifolia 61 West Heslerton Brow Good, R. D’O. 1956 Galeopsis angustifolia 62 Ellerburn Worsdale, A. F. 1952 Clinopodium menthifolium 10 Apesdown Stratton, F. 1872 Clinopodium menthifolium 10 Tucker, R. 1875 Clinopodium menthifolium 10 Apesdown Melvill, J. C. 1882 Galium tricornutum 9 Portland Good, R. D’O. 1919 Galium tricornutum 19 Saffron Walden Groves, J. 1875 Galium tricornutum 37 Craycombe Hill Baxter, T. 1860 Galium tricornutum 37 Upper Howsell, Malvern Baxter, T. 1860 Galium tricornutum 38 Whitnash 1879 Centaurea calcitrapa 13 Brighton 1872 Centaurea calcitrapa 15 Shoreham 1881 Centaurea calcitrapa 17 Barnes Groves, J. 1874 Centaurea calcitrapa MS ee ersey; Piquet, W. 1878 Hieracium pseudocurvatum 88 Killin Linton, W. R. 1883 Hieracium pseudocurvatum 92 Ben na Mac Duhi 1846 Hieracium pseudocurvatum 92 Corrie Etachen Linton, E. F. 1884 Hieracium pseudocurvatum 92 L. Kander 1885 Hieracium pseudocurvatum 108 Ben Loaghal Shoolbred, W. A. 1897 and Marshall, E. S. Pulicaria vulgaris 9 Portland Island - Pulicaria vulgaris 37 Malvern Baxtermul: 1857 Damasonium alisma 17 Thames Ditton 1874 Damasonium alisma 17 Mitcham Common Bennett, A. 1881 Potamogeton acutifolius 14 Pevensey Levels Wilson, A. K. 1934 Schoenoplectus triqueter 17 Bank of Thames Newnham, W. O. 1846 Schoenoplectus triqueter 17 Putney, by the Thames Bennett, A. 1874 Schoenoplectus triqueter 17 Mortlake Groves, H. and J. 1875 Schoenoplectus triqueter 17 Kew - Richmond Morgan, J. H. 1893 Lolium temulentum 1 Penzance 1878 Lolium temulentum 3 Plympton ' Briggs, T. R. A. 1878 Lolium temulentum 59 Liverpool, north of Lewis, J. H. 1872 Cypripedium calceolus 64 Grassington H I for F A L [Lees] 1878 HLL critically endangered taxa Ranunculus arvensis 61 King George Dock, Hull Wilson, A. K. O37 Scandix pecten-veneris 61 Cottingham 1864 Scandix pecten-veneris 61 King George Dock, Hull Wilson, A. K. NB Galeopsis angustifolia 63 Kirk Smeaton Rob, C. M. O37) Galium tricornutum 61 King George Dock, Hull Wilson, A. K. N37) Centaurea calcitrapa 41 Splott, Cardiff Smith, R. L. 1937 Potamogeton acutifolius 14 Pevensey Levels Wallace, E. C. 1934 Lolium temulentum 41 Cardiff Smith, R. L. 1938 runs, with little attention, on a PC running the Microsoft Internet Information Server (IIS) software. Although this system may not be suitable for servicing a large number of requests, it is more than adequate for the level - of use experienced and expected. The response to the pilot launch was both exhilarating and disheartening. It was clear that there were many specimens in the herbarium that were of interest to others and that a valuable service could be provided by putting the catalogue into the public domain. Feedback THE UNIVERSITY OF HULL HERBARIUM 337 from users did, however, reveal the shortcomings of the index. Many data entry errors were identified, mainly involving the spelling of localities and assignment of v.c. numbers. Subsequent programmes of checking have considerably reduced these errors. More importantly, feedback has enabled corrections and additions to be made on the basis of information provided by others and _ not available when the catalogue was compiled. In addition to the internet implementation of the catalogue, a stand-alone version has also been produced on compact disc (CD). This consists of a dedicated program and a_ specially formatted dataset. With this version it is possible to perform more _ sophisticated searches, there is no need for an internet link and it has been possible to integrate scanned documents into the database. The major disadvantage is that the program is platform specific (PC only) and the database is fossilised. Corrections and updates can easily be applied to the internet version but require a new version of the CD. Despite the drawbacks it is considered that the CD version will probably be the most convenient for persistent users and the internet version better for casual use. The system of dissemination described here is flexible and easy to implement. Although unsuitable for large herbaria, it would certainly be adequate for collections of up to 100,000 specimens. The most time-consuming element of the process has been the data entry and validation and any herbarium already having a computerised catalogue could implement a similar system with a relatively small invest- ment. The ready availability and low cost of modern computers now opens up even more radical prospects. It would be a relatively simple matter to scan the sheets and make the collection available as a virtual herbarium. Although this would not obviate the examin- ation of the specimens for some purposes, most users would probably not need to use the originals. In addition to saving wear and tear on fragile specimens it would open up access to the herbarium by making it universally available. ACKNOWLEDGMENTS The authors are grateful to The Feren’s Trust, University of Hull and The Botanical Society of the British Isles for grants which have enabled this work to proceed. Thanks are also due to Prof. David Taylor, now at Trinity College, Dublin, who originally instigated the cataloguing and to Mark Anderson, Nigel Arnett, Chris Blythe, Marion Brazier and Alan Drummond who have worked on the database at various times. Thanks are also due to Dr R. Goulder for his helpful comments on_ the manuscript. REFERENCES ANON. (1878). Barrow Naturalists’ Field Club 1877-1878. Proceedings of the Barrow Naturalists’ Field Club and Literary and Scientific Association 2: 4-7. ANON. (1902). Botany Report. Proceedings of the Barrow Naturalists’ Field Club and Literary and Scientific Association 14: 4—7. ARNOLD-FORSTER, K. (1993). Held in Trust. HMSO, London. BAMFORD, T. W. (1978). The University of Hull : the first fifty years. Oxon University Press, Oxford. BRITTEN, J. & BOULGER, G. S. (1931). A biographical index of deceased British and Irish botanists. (revised and completed by A. B. Rendle) Taylor & Francis, London. CHRISTIAN, W. H. (1935). Botanical Report. Proceedings of the Barrow Naturalists’ Field Club and Literary and Scientific Association 3(New series): 25-26. CHRISTIAN, W. H. & RICHARDSON, F. N. (1933). Botanical Report. Proceedings of the Barrow Naturalists’ Field Club and Literary and Scientific Association 2(New series): 57-58. CLEMENT, E. J. & FOSTER, M. C. (1994). Alien plants of the British Isles. Botanical Society of the British Isles, London. CRACKLES, F. E. (1990). Flora of the East Riding of Yorkshire, Hull University Press & Humberside County Council, Hull. DANDY, J. E. (1958). List of British vascular plants. British Museum & Botanical Society of the British Isles, London. DESMOND, R. (1994). Dictionary of British and Irish Botanists and Horticulturalists. Taylor and Francis, London. EDEES, E. S. (1972). Flora of Staffordshire. David and Charles, Newton Abbot. Goop, R. (1948). A geographical handbook of the Dorset Flora. The Dorset Natural History and Archaeological Society, Dorchester. 338 R. MIDDLETON & J. R. MIDDLETON HULL UNIVERSITY ARCHIVES, Ronald Good’s Letters. DX/14/4 KENT, D. H. (1992). List of vascular plants of the British Isles. Botanical Society of the British Isles, London. KENT, D. H. & ALLEN, D. E. (1984). British and Irish Herbaria. Botanical Society of the British Isles, London. RYVES, T. B., CLEMENT, E. J. & FOSTER, M. C. (1996). Alien grasses of the British Isles. Botanical Society of the British Isles, London. SLEDGE, W. A. (1975). Catherine Muriel Rob F.L.S. Naturalist 100: 67-68. VENN, J. A. (1954). Alumni Cantabrigienses. Cambridge University Press, London. WALSH, G. B. (1944). Thomas Stainforth BA BSc (1882-1944), in memoriam. Naturalist 69: 77—78 WILSON, A. K. (1938). The Adventive Flora of the East Riding of Yorkshire. Occasional papers of the Hull Scientific and Field Naturalists’ Club 1, Hull. (Accepted August 2006) Watsonia 26: 339-345 (2007) 309 A new far northern hybrid horsetail from Scotland: Equisetum xmchaffieae C. N. Page (Equisetum fluviatile L. x E. pratense Ehrh.) CN: PAGE* Gillywood Cottage, Trebost Lane, St. Stythians, Truro, Cornwall TR3 7DW H. McHAFFIE Royal Botanic Garden, Edinburgh EH3 5LR J. K. BUTLER Seaside Cottage, Thurso East, Caithness KW14 SHN ABSTRACT A new hybrid horsetail Equisetum xmchaffieae C.N. Page hybr. nov., with the parentage E. fluviatile L. x E. pratense Ehrh., is described from Caithness, Scotland, v.c.109. It is only the second known British Equisetum hybrid involving the northern EF. pratense in its parentage, a species which itself is rare in the British Isles as a whole. A description is given, diagnosis with other hybrids made and the ecology of this unusual taxon outlined. KEYWORDS: Equisetum, hybrid, morphology, ecology, Caithness, Scotland. INTRODUCTION A new hybrid horsetail, Equisetum xmchaffieae C. N. Page, hybr. nov., with the parentage E. fluviatile L. x E. pratense Ehrh., is described. First found by Heather McHaffie in Caithness, v.c.109, northern Scotland, in August 2003 and submitted to the referee for Equisetum, this plant is clearly of hybrid origin and presents a distinctive individual vegetative morphology closely intermediate between that of its two, somewhat contrasting, parents. It is only the second known British Equisetum hybrid in- volving the northern E. pratense in its parentage, a species which itself is scarce in the British Isles as a whole. The formal description given here follows the initial format of most other hybrids in the Equisetum subgenus Equisetum (e.g. Page 1963, 1973, 1988a, 1995; Dines & Bonner 2002) to facilitate comparison. Further morphological details are given, diagnosis with other hybrids in Egquisetum subgenus Equisetum involving either parent made, and the ecology so-far known of this unusual taxon outlined. *e-mail: pterido@hotmail.com DESCRIPTION AND DIAGNOSIS Equisetum xmchaffieae hybr. nov. (E. fluviatile L. x E. pratense Ehrh.) Fig. 1 Caules 25-45 cm alti, 1-8—2-3 mm diameter, viridis, alte 7—12 sulcati, porci angulosus, pagina minutus asperatus; vaginae (dentibus exclusis) 3:5—4-1 mm, virides; dentes tot quot sulci, subulati, piliferous-acuminatus, nigres- centes, marginibus basium angustis scariosis, l-costati; cavitas centralis 1/2 caulis diam. Ramuli patentes vel suberecti, regularites verticillati, simplices, tenuis, plerumque tri- gonis et tetragonis, carina aculis, internodium infimum vaginam caulis proximam aequans vel 1-5 superans; vaginae pallidae, dentibus acuminatis, patens, viridis. Stems 25-45 cm, erect, 1-8—2:3 mm diameter, green, grooves 7-12, ridges angular, surface minutely rough; sheaths (excluding teeth) 3-5- 4-1 mm, green; teeth as many as the grooves, subulate, piliferous-acuminate, blackish with narrow scarious basal margins, 1-ribbed; central hollow 1/2 diameter of stem. Branches spreading to suberect, in regular whorls, simple, slender, three or four angled, ridges acute, the lowest internode 1:0—1-:5 times the length of the adjacent stem sheath; sheaths pale, teeth acuminate, non-spreading, pale green. HOLOTYPUS: Scotland, Caithness, St. John’s Loch, Scotland, U.K., GR ND 220 718, 12 August 2003, Heather McHaffie (E). Equisetum xmchaffieae is clearly intermediate in shoot vegetative structure between E. fluviatile and E. pratense. Plants have slightly C. N. PAGE, H. McHAFFIE & J. K. BUTLER 340 = ) —] — FIGURE 1. Silhouette of type material of Equisetum xmchaffieae C. N. Page, hybr. nov. (E. fluviatile L. x E. pratense Ehrh.). EQUISETUM xMcHAFFIEAE 341 decumbent to erect main stems 25—45 cm in height, but only 1-8—2:3 mm shoot diameter, with a broadly ovate outline closely remin- iscent of that of E. xlitorale (E. arvense x fluviatile), but with overall more slender, more delicate structure throughout and with more numerous notably more slender branches. Nodes throughout the middle 3/4 of the shoot are branch bearing, with typically 3-4 or more at the base of the shoot non-branch-bearing and 5—6 or more of the most distal forming a long slender 0-6—0-8 mm diameter or less gradually tapering terminal ‘tail’ to the upper extremity of each shoot. Main shoot internodes with numerous (7-12) shallow, highly angular ridges and grooves, with an overall minutely roughened surface texture of fine siliceous spicules detectable especially when a finger nail tip is drawn gently along. All main stem internodes also lack the easy elastic com- pressability in fresh material of those of E. xlitorale resulting from the central canal of this hybrid being mostly little more than 1/2 the overall diameter of the stem combined with smaller diameter and tougher shoot structure. Characteristic main stem sheaths throughout the lower third of the shoot are scarcely inflated but strongly vertically grooved, c. 2:4 mm diameter and 3:-5—4:0 mm long excluding teeth, green throughout or sometimes irregularly dark-patched at the base, their free teeth particularly numerous (7-12) for the sheath width, up to 1-9 mm long, dark brown to blackish throughout, initially broad at the base but tapering rapidly with narrow pale brown scarious margins in the basal portions only, and for their final half to two thirds (c.0-8—1-0+ mm) tapering only slowly to form distinctive, individual, relatively long, straight, narrow, fine, entirely black, somewhat delicate piliferous acuminate teeth tips, which can become rather easily broken free in dried material. Branches slender and _ particularly numerous, up to 12 per whorl, neatly regi- mented, spreading-ascending to somewhat upward curving throughout the shoot, c. 9:5— 10-5 cm long in the middle of the shoot, reducing to 4:5—5:5 cm in the upper part of the shoot, c. 0-6 mm or less diameter, their inter- nodes (3)-4 angled, strongly furrowed, the furrow base with a shallow sulcus, the ridges acute, minutely pearled along their apices but not saw-toothed, the branch sheaths green, uniangulate, the teeth simple, mostly pale green sometimes tinged translucent brown towards the tips, slender, straight, not spreading. The hybrid is named after Heather McHaffie, who collected the material, first recognised it to be distinctive in the field, and submitted it to the referee: TAXONOMIC COMPARISONS Equisetum xmchaffieae is a distinctive hybrid, with shoots which are strikingly intermediate in size, morphology and anatomy between the somewhat contrasting morphologies of E. fluviatile L. and E. pratense Ehrh. As seems characteristic of a majority of colonies of hybrid horsetails, and often a feature of them, no shoots of this hybrid have yet been found to bear cones, but their strong intermediacy of vegetative structure is sufficiently consistent and distinctive to be a reliable guide to parentage, with clear characters which link directly with each of the above species. Equisetum xmchaffieae possesses the overall markedly ovate E. fluviatile-dominated overall shoot outline characteristic of all of the known hybrids of this parent (EF. x/itorale Kuhlew. ex Rupr., E. xdycei C. N. Page, E. xwillmotii C. N. Page), and especially with a branchless portion at the base and a relatively long branchless terminating ‘tail’ portion to the tip of the shoot. However E. xmchaffieae has an overall marked slenderness and delicacy of form with a thin but tough and angular main stem and overall slender branch structure readily apparent both in the field and in herbarium material, which distinguish it especially from the above hybrids. Further, the distinctive minute roughness of the main stem internodes to touch is unusual for any hybrid and is characteristic only of hybrids involving either E. pratense or E. sylvaticum in their parentage, but especially the former, which the notable angularity of the ridges in its main shoot cross-section also most closely appr- oaches. Additionally in E. xmchaffieae, the numerous long, straight, narrow, fine, delicate, black, entirely free teeth to the main stem sheaths through at least the lower third of the main shoot, with markedly piliferous acuminate tips, the highly angular slender branches, and lack of secondary branching of the branch systems, all confirm the unusual involvement of E. pratense in the parentage of this hybrid. In more detailed comparisons with the other Equisetum fluviatile hybrids, Equisetum xmchaffieae is thus superficially similar mainly 342 C. N. PAGE, H. McHAFFIE & J. K. BUTLER only to E. xlitorale in general outline, in comparison with which it looks smaller, much more slender and delicate, with clearly rougher surface texture, longer, narrower, numerous, dark, free, main stem teeth, and lack of the the ‘plastic-like’ yield (Page 1997a) of the main shoot internodes when gently squeezed between finger and thumb, so characteristic in the field of E. x/itorale. In comparison with E. xdycei, E. xmchaffieae has also a typically rougher surface and more copious spreading- ascending more angular, more slender, bran- ches and the distinctive E. pratense-like teeth to the main stem sheaths; and in comparison with E. xwillmotii is also of very much more overall slender and more delicate structure, with rougher main stem internode surfaces and more slender, simple, uniangulate, branch internodes with sheaths which are _ not bicarinate. In more detailed comparison with E. xmildeanum, the only other Equisetum pratense hybrid so-far known in Britain, E. xmchaffieae differs clearly in possessing only simple (ie. unbranched) branch structure and in possession of the distinctive terminal ‘tail’ to the tip of the shoot, more often 4-angled branches (combining the tendency of respective parents to be dominated by 3 and 4—5 angled branches respectively), and clearly simpler more separated free teeth segments to the main shoot sheaths. However, this particular hybrid would most easily be dismissed (as might also E. xdycei) in the field as merely a slender and perhaps depauperate form of E. x/litorale (as might also E. xwillmotii, for example, as being a more robust form of E. xlitorale), and thus either mis-recorded or overlooked or not collected at all for being ‘aberrant’. It may well be that the occurrence of this hybrid has been overlooked for these reasons. However, this hybrid is between two species of Egquisetum which would also appear to not be especially closely- related (Page 1972), and the phylogenetic distance apart of their parents (but within the same subgenus) would seem likely to be a factor in its apparent rarity of occurrence. This distance apart, however, approximates also to that of the parents of each of E. xwillmotii, E. xbowmanii and E. xrobertsii, all of which have so-far proved to have a relatively small number of scattered stations in Britain and Ireland (in comparison with the relative frequency of E. xlitorale sites — now well over 100 in these islands), and all too, typically not necessarily always with both parents present (an aspect. which may well echo past distributions, which the hybrids have outlived). FIEED ECOLOGY Equisetum xmchaffieae occurs at the edge of St. John’s Loch, Caithness, where it is associated with a slightly flushed fen near to the water’s edge. The surrounding overall vegetation type includes mainly Filipendula ulmaria, Potentilla palustris, Salix repens, Eriophorum angustifolium, Succisa pratensis, Achillea ptarmica, Senecio palustris, Parnassia palustris, Caltha palustris, Triglochin palustris, Lychnis flos-cuculi, Hydrocotyle vulgaris, Stachys palustris, Juncus effusus, J. conglomeratus, Carex panicea, Calamagrostis stricta, Molinia caerulea, Holcus lanatus, Anthoxanthum odoratum, Deschampsia cespi- tosa, Equisetum fluviatile and Equisetum palustre. There are strips of fields running between a nearby road and the water’s edge, - with the upper part of each field above the 25 m contour usually cultivated, but their lower edges at least 10 m wide towards the loch are not cultivated, though might be sometimes grazed. The loch is freshwater, with a water level artificially maintained for native brown trout, for which there is a hatchery, and a track to the water’s edge to access fishing boats. Of the parent species, Equisetum fluviatile is present around the loch margins. E. pratense is not present immediately nearby, but is however, not insignificant in occurrence around the overall vice-county and adjacent region and has long been known from both 10 km grid squares immediately adjacent to the south of the loch (where it is most likely present along banks and sandy shoulders of upland streams and rivers feeding the lochs). It also grows as a substantial colony on Thurso riverside at ND1113673, and has also prev- iously been found in rough vegetation at Dunnet links (ND215688, 1997, although its present survival at this location is questioned). Within this overall setting, Equisetum xmchaffieae 1s present in a small bay at the edge of the loch over a substratum of shelly clay originally of marine origin deposited within an area otherwise bounded by Old Red Sandstone. In terms of micro-topography, the hybrid occurs clinging to the wave-worn bare vertical clay created by wave action and through the vegetation behind. This sheer bank is some | m high topped by a thin layer of peat and bears the same flora as above. At the loch, hybrids of Equisetum would appear to be relatively vigorous, since, other collections from this location recognised as unusual in the field by Heather McHaffie and submitted to the referee for checking have proved to contain EQUISETUM xMcHAFFIEAE 343 also E. xlitorale, E. xdycei and E. xrothmaleri at this site. Shoots of what are possibly all of E. xmchaffieae are scattered at a frequency of maybe | shoot per metre over a length of | km of shore explored, centred on the grid point ND22007211, and with a possible higher den- sity in parts. Overall, members of the genus and especially Equisetum hybrids as a whole are clearly remarkably successful in this overall location, and their patterns of distribution are likely complex and may well be highly intermixed. DISCUSSION Equisetum xmchaffieae is the sixth known hybrid in the genus Equisetum recognised within Scotland, and brings to a total of eleven the number of inter-specific hybrids within the genus as a whole known within Britain and Ireland. It also brings to nine, the number of hybrids known within Equisetum subgenus Equisetum (the deciduous-shooted horsetails) formed between the six native species of this subgenus in these islands. Eguisetum xmchaffieae is also the fourth known hybrid to involve E. fluviatile L. in its parentage: the others being [in order of frequency] E. xlitorale (E. arvense x fluviatile), E. xdycei (E. fluviatile x palustre) and E. xwillmotii (E. fluviatile x telmateia), all of which hybrids are known in Britain now in multiple locations. Equisetum xmchaffieae is, however, only the second known hybrid in Britain to involve E. pratense in its parentage, the other being E. xmildeanum Rothm. (E. pratense x sylv- aticum), which has now _ several known scattered stations in highland Scotland and in the mountains of northern and central Europe (Rothmaler 1944; Page 1988a, 1997a; Lubien- ski 2003). No known extra-British localities for this particular hybrid have, however, been either reported or as yet found in herbarium material from elsewhere, although, of its parents, both are species with extraordinarily wide temperate ranges (Hultén & Fries, 1986). Equisetum fluviatile is circumboreal, occur- ing across Eurasia and far eastern Asia to Japan, northward from approximately latitude 36 degrees N and across North America at similar latitudes (Hultén & Fries 1986; Derrick et al. 1987). In Europe, E. fluviatile occurs widely from extreme southern Greenland, Iceland, and northern Scandinavia to approxi- mately the Pyrenees and Caucasus, with outlying stations to southern Spain and Italy (Jalas & Suominen 1972). It is most abundant at middle to northerly latitudes, where it is a species of “shores of lakes and slow rivers, often bordering reed swamp communities down to 1:5 m depth, forming large pure stands especially in oligotrophic lakes, mesotropic and eutrophic mires, margins of bogs and oligotrophic fens, Salix and Alnus carrs” (Jonsell 2000). Within Britain and Ireland, E. fluviatile is similarly widespread, from the Orkneys and Shetlands to Cornwall @enny wen ala N97S Preston ) er malo 2002); especially at low to moderate elevations, and of a similar habitat range, but mainly a species of low to moderate altitude, part-shaded to exposed, lake and other of standing water margin, ditch and carr vegetation communities, sometimes forming dense and extensive cone- bearing stands in open water reedswamp communities. In Scotland it occurs quite widely at lower altitudes but ascends as mostly smaller-statured, often cone-inhibited, sparser stands to alpine lochans and even into wet heathland communities to an altitude of about 900 m. Equisetum pratense is an _ arctic-alpine species which is most abundantly northern in its circumboreal range. It is present across Eurasia to the Altai, Manchuria, Kamtchatka and the Chukotsk Peninsula of far eastern Siberia, most abundantly north of approx- imately latitude 48 degrees N, and at similar latitudes across North America from Alaska to Labrador (Rothmaler 1944; Lellinger 1985; Hultén & Fries 1986; Derrick et al 1987). In Russia, it is widespread and locally frequent across northern Russia and especially in Siberia from the South Siberian mountains northward (Komarov 1934; Krasnoborov 1988); and in northwestern Canada sometimes forms ‘dense stands on open wooded floodplains of rivers and streams’ (Cody & Britton 1989). In western Europe, it ranges from Iceland and northern Scandinavia to Central Europe and the Alps (Jalas & Suominen 1972; Hegi 1984), and especially in more northerly latitudes can be locally com-mon mainly on moist, base rich substrates in open woods, pastures, river banks, hay-meadows, alpine tall-herb communities, snow-beds and talus slopes (Jonsell 2000). Within Britain and Ireland, E. pratense is confined to the northern half of these islands, where it has scattered stations in the northern Pennines and in Northern Ireland, but more widely spread only in Scotland and especially in the central and eastern Highlands and north 344 C. N. PAGE, H. McHAFFIE & J. K. BUTLER to Caithness (Jermy et al 1978, 1994; Preston et al. 2002). In this range, E. pratense is a scarce British species largely of sandy-clayey river and streambanks, especially in moist, lightly shaded slopes, hence chiefly in the lower valleys of mountain districts, in sites typically where the ground contains a high mineral component, and this is especially of sand which is either calciferously derived or is base-flushed. In Scotland it ascends as more scattered populations near to tumbling mount- ain rills and on ledges over mica-schist rock to at least 915 m (and reported to ascend to 1,400 m in the mountains of mainland Europe (Prelli 2001). It is, however, only in its more northern stations in Britain (central Perthshire north) that I have found E. pratense currently to produce fertile cones (though a century ago, herbarium evidence shows that colonies were freely fertile as far south as Coquetdale, Weardale and Teesdale (Page 1988b, 1997b), where recent fieldwork shows the plant to still persist, but in entirely vegetative condition. Equisetum. xmchaffieae thus occurs within the northern part of the sympatric portion of the ranges of its two parents in Britain, and especially within the zone in which both are cone-bearing. The geological aspects of its Caithness site are also clearly especially suited to Equisetum growth as a whole, occurring within an area bounded by Old Red Sandstone and directly over a substrate of shelly clay of originally marine origin, overlain by peat and now exposed in a small freshwater lochside bay. For such exposed clay is a particularly edaphically-appropriate mineral habitat for the occurrence of species of Equisetum, since it provides the essential mineral combination of a substrate combining both available silica and calcium at the same time, here the calcium- source being especially in the form of the marine shell component within the sandy-clay matrix (Page 1988b, 1997). The close association of this hybrid in this location with the face of a | m high eroding bank face of this shelly clay also appears significant. Clearly this face must afford rather better drainage than would occur within the fen vegetation itself, but nevertheless be influenced by both moving ground moisture and locally enhanced hum- idity. Further, the association of this hybrid with such a clear erosion-surface is notable as such erosion-surfaces which are in a more or less constant state of micro-disturbance are believed to be especially important in prov- iding sites for the co-occurrence of prothalli of disparate species of Equisetum within cross- fertilisation distance, and hence especially effective in the induction of hybrid formation, in climates as recurrently moist and relatively mild as many of those of the British Isles, where, by European standards as a whole, hybrids within Equisetum subgenus Equisetum are especially rich (Page 1979, 2002; Page & Barker 1985; Page & McHaffie 1991; Dines & Bonner 2002). Here, at this Caithness site, the presence of such a concentration of co-extent different Equisetum hybrids (E. xlitorale, E. xdycei, E. xrothmaleri and E. xmchaffieae), collectively involving different combinations of four local parent species, emphasises the significance of the combined role of appro- priately moist climate, edaphic mineral balance, exposure of the mineralised surfaces and recurrent small disturbance processes operating on these, can generate in opening the opportunity for hybrids in this genus to form. ACKNOWLEDGMENTS The authors are grateful to Professor Mary Gibby, Douglas McKean and Dr Robert Mill (RBG Edinburgh) for helpful comments on the manuscript. REFERENCES Copy, W. J. & BRITTON, D. M. (1989). Ferns and Fern Allies of Canada. Agriculture Canada, Ottowa. DERRICK, L. N., JERMy, A. C. & PAUL, A. M. (1987). Checklist of European pteridophytes. Sommerfeltia 6: 1-94. DINES, T. D. & BONNER, I. R. (2002). A new hybrid horetail, Eguisetum arvense x E. telmateia (E. xrobertsii) in Britain. Watsonia 24: 145-157. HEGI, G. (1984). Ilustrierte Flora von Mittel-Europa. Band I. Pteridophyta. Paul Parey, Berlin & Hamburg. HULTEN, E. & FRIES, M. (1986). Atlas of North European Vascular Plants. Konigstein, Federal Republic of Germany. JALAS, J & SUOMINEN, J (1972). Atlas Florae Europaeae. |. Pteridophyta. Committee for mapping the Flora of Europe, Helsinki. JERMY, A. C. (1994). Equisetum pratense Ehrh., in A. STEWART, D. A. PEARMAN, & C. D. PRESTON. eds. Scarce Plants in Britain, p.154. Joint Nature Conservation Committee, Peterborough. EQUISETUM xMcHAFFIEAE 345 JERMY, A. C., ARNOLD, H. R., FARRELL, L. & PERRING, F. H. (1978). Atlas of Ferns of the British Isles. Botanical Society of the British Isles & British Pteridological Society, London. JONSELL, B. (2000). Flora Nordica. The Royal Swedish Academy of Sciences, Stockholm. KOMAROV, V. L. (ed.). (1934). Flora of USSR. Vol. 1. Academy of Sciences of USSR, Leningrad. KRASNOBOROY, I. M. (ed.). 1988. Flora of Siberia. Vol. 1. Nauka, Novosibirsk. LELLINGER, D. B. (1985). A Field manual of the Ferns and Fern-allies of the United States and Canada. Smithsonian Institute Press, Washington, D.C. LUBIENSKY, M. (2003). Equisetum xmildeanum Rothm. (E. pratense Ehrh. xE. sylvaticum L.), en snellehybrid ny for Norge. Blyttia 61: 171-178. PAGE, C. N. (1963). A hybrid horsetail from the Hebrides. British Fern Gazette 9: 117-119. PAGE, C. N. (1967). Sporelings of Equisetum arvense in the wild. British Fern Gazette 9: 335-338 PAGE, C. N. (i972). An assessment of inter-specific relationships in Eguisetum subgenus Equisetum. New Phytologist 71: 355-369. PAGE, C. N. (1973). Two hybrids in Equisetum new to the British Flora. Watsonia 9: 229-237. PAGE, C. N. (1979). Equisetum xtrachyodon in western Scotland. Fern Gazette 12: 178-179 PAGE, C. N. (1988a). Two hybrids of Eguisetum sylvaticum new to the British Flora. Watsonia 17: 2273-277. PAGE, C. N. (1988b). Ferns: Their Habitats in the Landscape of Britain and Ireland. Collins New Naturalist no 74, London & Glasgow. PAGE, C. N. (1995). Equisetum xwillmotii C. N. Page — a new hybrid horsetail from County Cavan, Ireland. Glasra 2: 135-138. PAGE, C. N. (1997a). The Ferns of Britain and Ireland. 2nd edn. Cambridge University Press, Cambridge. PAGE, C. N. (1997b). Pteridophytes as field indicators of natural biodiversity restoration in the Scottish flora. Botanical Journal of Scotland 49: 405-414. PAGE, C. N. (2001). Ferns and allied plants, in D. L. HAWKSWORTH ed. The changing wildlife of Great Britain and Ireland. Systematics Asociation Special Volume 62, pp 50-77. Taylor & Francis, London & New York. PAGE, C. N. (2002). The role of natural disturbance regimes in pteridophyte conservation management. Fern Gazette 16: 284-290. PAGE, C. N. & BARKER, M. A. (1985). Ecology and geography of hybridisation in British and Irish horsetails. Proceedings of the Royal Society of Edinburgh 86B: 265-272. PAGE, C. N. & MCHAFFIE, H. S. (1991). Pteridophytes a indicators of landscape changes in the British isles in the last hundred years, in J. M. CAMUS ed. The History of British Pteridology, pp. 25-40. British Pteridological Society, London. PRELLI, R. (2001). Les Fougeres et Plantes Alliees de France et d’Europe Occidentale. Belin, Paris. PRESTON, C. D., PEARMAN, D. A., & DINES, T. D. (2002). New Atlas of the British and Irish Flora. Oxford University Press, Oxford. ROTHMALER, W. (1944). Pteridophyten Studien I. Feddes Repertorium 54: 55-82. (Accepted June 2006) | ue | m7 : i ‘ ho : _ 7 eal ee eee | i on a : 7 “i il a = i . as i + y i. a4 _o i : a U 7 , = i ‘ * cc 2 1 r , i) . | Watsonia 26: 347-358 (2007) 347 A fine scale study of selected environmental and floristic parameters in three populations of Euphrasia vigursii (Davey), a rare annual endemic to Devon and Cornwall L. GRANADOS' and S. D. LANE” School of Biological Science, University of Plymouth, Plymouth PL4 8AA ABSTRACT Euphrasia vigursii is an endangered hemiparasitic annual, endemic in Devon and Cornwall. This investigation has evaluated fine scale patterns of floristic composition and nutrient distribution in selected populations of FE. vigursii in Devon and Cornwall to increase understanding of their ecology for use in future management strategies in Devon. Two particular questions were formulated: 1. Is E. vigursii immediately surrounded by other particular plant species? 2. If the soil was analysed in the direct location of the site of E. vigursii would the pH, Nitrogen, Phosphorus and Potassium content be different from the areas where E. vigursii is not present? Phytosociological analysis indicated that E. vigursii is associated with heath and acidic grassland com- munities typically containing Ulex gallii, Agrostis curtisii, Calluna vulgaris, together with Molinia caerulea and Potentilla erecta mire. Within the individual sites spatial scale comparison (macro vs. micro) indicated significant differences in Phosphorus levels and pH levels but these were not consistent across the sites. Germination of E. vigursii at microenvironment level was closely correlated with the presence of A. curtisii rather than its other known host species Ulex gallii. This is contrary to published research which suggests that E. vigursii avoids the more open, species-poor areas dominated by A. curtisii. Correlations between the environ- mental parameters assessed may reflect present man- agement regimes within the Dartmoor populations. KEYWORDS: spatial scale, species distribution, hemiparasitic, soil nutrients, conservation, Vigurs’ Eyebright. INTRODUCTION E. vigursii (Vigurs Eyebright) is confined to Cornwall and Devon where it is considered endangered and is classified as vulnerable (Cheffings & Farrell 2005). Until 1979 it had been located in 53 sites in Cornwall; later 'e-mail:lisette.granados @ntlworld.com 2e-mail: slane@ plymouth.ac.uk surveys have revealed only 25 active sites. This represents approximately 50% loss of localities in Cornwall mainly due to habitat destruction (French et al. 1999). In Devon there are only 2 sites, rediscovered in 1995 and 1998 and situated on the north-west perimeter of Dartmoor National Park. Ecologists at the National Park aim to conserve and increase the population of E. vigursii as part of the Dartmoor Biodiversity Action Plan, as it has been categorised in the priority list of the UK steering group report on biodiversity as being of ‘global conservation concern’ (HMSO 1995). In order to establish a management strategy that would address the objectives of Dartmoor National Park authority, a greater understanding of the ecology of E. vigursii and specifically knowledge of the local populations is essential. The processes and mechanisms that result in species being rare and endemic are complex traits involving interactions of abiotic, biotic and historical factors. Investigating the factors which influence the distribution of E. vigursii must therefore involve an amalgamation of demographic, genetic, resource allocation Wavy vec Setlenesss 198i; Palmer, 1987: Schemske ef al., 1974) alongside historical systematic and stochastic perturbations (Shaffer 1981). E. vigursii is a hemiparasitic summer annual, an erect to 20 cm branched therophyte (Grime 1996) located in heathland containing Ulex gallii and Agrostis curtisii in both Devon and Cornwall. The leaves are dull greyish green often suffused with violet or black (Stace 1999) due to anthocyanin pigment (Pankhurst 1977). The corolla is usually lilac/purple/reddish and on the upper leaves and bracts are long glandular hairs. E. vigursii is only one of four diploids in the genus and considered a stable hybrid between Euphrasia micrantha and 348 L. GRANADOS & S. D. LANE ‘ Somerset Devon And Dorset Site 1: Lydford Dartmoor \ = N Site 2: St Neote iy , ci = W E Site 3: St Agnes Cornwall_,~} eo x : < Scale: 16 km FIGURE |. Location of study sites in Devon and Cornwall. Euphrasia anglica (Pankhurst 1977). An open grass sward provided by regular grazing and controlled burning (swaling) is considered characteristic of the habitat in which E. vigursii is found (Norman Baldock, Ecologist Dartmoor National Park; pers. comm.. 2000). Very little specific research on E. vigursii has been con- ducted: therefore, the aims of this study have been to evaluate some of the ecological and environmental parameters influencing three separate populations of E. vigursii to gain better understanding of its ecology and habitat requirements and to answer two _ specific questions: 1. Is E. vigursii immediately surrounded by other particular plant species? 2. If the soil was analysed in the direct location of E. vigursii would the pH, Nitrogen, Phosphorus and Potassium con- tent differ from areas where E. vigursii is not present? MATERIALS AND METHODS Two surveys were carried out at each of three locations in Devon and Cornwall (Fig. 1). The first survey entailed micro sampling of the target species FE. vigursii to determine at individual plant level the composition of neighbouring plant associations together with recordings of soil nutrients (Potassium, Phos- phorus and Nitrogen) and pH status. The second survey involved macrosampling to give a general indication of the vegetative com- munity composition in which E. vigursii was found together with the soil nutrient and pH status within areas where E. vigursii was not present. SITE 1 Lydford, Dartmoor National Park, Devon. Altitude: 320 m. Sample site size: 10 m x 50 m In the surveyed area Ulex spp. (gorse) and heather formed a mosaic of islands surrounded by pathways containing a grass and herbaceous sward. E. vigursii was located in most instances at the parameter of mature and new vegetative growth of U. gallii and where A. curtisii in the ‘pathways’ was of low sward height and also where periodic swaling had taken place. SITE 2 St. Neot, Bodmin Moor Cornwall. Altitude: 170 m. Sample site size: 7 m x 50 m. The surveyed area contained a linear grass ECOLOGY OF EUPHRASIA VIGURSII 349 and herbaceous sward, 15 m wide, which cut through a patchy landscape of gorse, heather and scrubland at the perimeters. FE. vigursii was situated within the short grass sward from the edge of tall Ulex spp. at the perimeter of the survey area to within 5 m of the linear grass sward. SITE 3 St. Agnes, nr Redruth, Cornwall. Altitude: 85 m. Sample site size: 3-5 m x 50 m In the surveyed area there was a blanket of dwarf gorse and heather species with pathways formed by walkers and erosion by the weather. E. vigursii was situated approximately 20 m from the cliff edge on one of the many worn paths that connect the coastal path that runs along the cliff edge with a parallel inland pathway. It was present amongst shorter veg- etation at each side of the pathway although a greater percentage of the population was situated north east facing and was therefore sheltered from south-westerly winds. SURVEY METHODS Each survey site was marked out into a 1 m? grid framework and random co-ordinates used for macro sampling (1 m° quadrats). Micro sampling used 40 mm circular quadrats that were positioned non-randomly; the quadrat was placed over the target species (E. vigursii in the centre) and species frequency within the microsampling area recorded. Soil samples were collected 10 mm from the stem of Euphrasia vigursii. For the macro survey a | m’ collapsible quadrat was randomly posi- tioned in the survey area and the percentage cover for each species was_ recorded. Identification of the floristic species was carried out using keys by Stace (1999), Hubbard (1985) and Fitter et al. (1995). Soil samples were taken from the centre of the quadrat using a 15 mm x 150 mm metal corer, to a depth of 25 mm to 75 mm de- pending on soil texture. In each survey site 60 soil samples were taken; 30 in the macro quadrats and 30 in the circular micro quadrats. ANALYSIS OF SOIL SAMPLES In total the surveys yielded 180 soil samples. Each was crushed and put through a 2 mm metal sieve before testing for pH, Potassium, Phosphorus, and Nitrogen using recommended methodology (ADAS Bulletin RB 427) for NPK and pH while C: N ratio was determined with a Leco FP 2000 C and N Analyser. COMMUNITY CLASSIFICATION AND PHY TOSOCIOLOGY In order to define the floristic community in which E. vigursii occurs the floristic data (using percentage cover data) were classified using two-way indicator species analysis and the computer program TWINSPAN. The TABLEFIT programme was then used to link the resulting communities in to the categories of the National Vegetation Classification (Dinsdale 1997). The total micro floristic data (frequency data) were also run through TWINSPAN and Detrended Correspondence Analysis computer program applied using PC ORD in order to examine which were the associative and most frequent floristic species with E. vigursii. This was carried out for the micro floristic data between and in each individual site. RESULTS Floristic data and soil cores were obtained from a total of 180 quadrats collected from the three sites. The 90 micro quadrats yielded 21 plant species in total while the macro quadrats contained 28 plant species (Table 1). Bare ground and dead material were also recorded . All the species occurred at both survey scales, apart from Poa spp. which was only found only in the micro quadrats. CLASSIFICATION AND PHYTOSOCIOLOGY Interpretation of two-way indicator species analysis and definition of plant communities The classification groups resulting from TWINSPAN analysis and synoptic frequency of occurrence within each group are shown in Table 2. Five TWINSPAN groups were defined and their associated NVC _ plant communities categorised (Table 3). Groups 1 and 2 were heath communities of Site 3, St Agnes. In group | Agrostis curtisii , Erica cinerea, Potentilla erecta, and Ulex gallii, Thymus spp., Serratula tinctoria and Ulex europaeus were frequent; however, there was a low frequency of Erica tetralix. In group 2 Ulex gallii, Thymus species, Erica tetralix, Carex flacca, Serratula tinctoria, Viola spp., Agrostis curtisii and Potentilla erecta were frequent. Dead material was a feature of both groups | and 2 and bare ground was a feature of the group 2. Group 3 were heath communities of Site 1, Lydford. Agrostis curtisti, Ulex gallii, Thymus spp., Potentilla erecta, Galium saxatile and 350 L. GRANADOS & S. D. LANE TABLE 1. ASSOCIATES OF EUPHRASIA VIGURSIT INMACRO AND MICRO SURVEY SITES IN DEVON AND CORNWALL Macro Micro Bare ground Dead material Agrostis capillaris Agrostis curtisil Anthoxanthum odoratum Calluna vulgaris Carex flacca Dactylis glomerata Danthonia decumbens Erica cinerea Erica tetralix Euphrasia anglica Euphrasia micrantha Euphrasia vigursii Galium saxatile Holcus lanatus Hypericum spp. Pilosella officinarum Plantago lanceolata Polygala serpyllifolia Potentilla erecta Pteridium aquilinum Rubus fruticosus Senecio jacobaea Serratula tinctoria Stachys officinalis Teucrium scorodonia Thymus spp. Ulex europaeus Ulex gallii Vaccinium myrtillus Viola spp. Agrostis capillaris Agrostis curtisii Carex flacca Erica cinerea Erica tetralix Euphrasia vigursii Galium saxatile Hypericum spp. Hypochaeris radicata Lotus corniculatus Pilosella officinarum Poa spp Potentilla erecta Senecio jacobaea Serratula tinctoria Hypochaeris radicata Lotus corniculatus Molinia caerulea Thymus spp. Ulex gallii Viola spp. Molinia caerulea were frequent but with low frequencies of Danthonia decumbens, Pilosella officinarum, Erica cinerea, Carex flacca and bare ground. Group 4 and 5 were a mixture of heath, mire, and under shrub communities of Site 2, St Neot. In group 4 Agrostis curtisii, Potentilla erecta, Pteridium aquilinum, Molinia caerulea, and Ulex europaeus were frequent with a low frequency but high abundance of Ulex gallii. Less common was Agrostis capillaris. In group 5 Agrostis curtisii, Potentilla erecta, Pteridium aquilinum, Agrostis capillaris, Senecio Jacobaea, Molinia caerulea and Galium saxatile and MHypochaeris radicata were frequent with Ulex gallii, Euphrasia vigursii, Carex flacca, Euphrasia anglica and Anthoxanthum odoratum less so. Euphrasia vigursii was found to be most frequent in group 2 (7 quadrats) at Site 3 compared to its presence in Site 3, group | (1 quadrat); Site 1, group 3 (1 quadrat) and Site 2, group 5 (4 quadrats). In group 2 it was associated with NVC U3, H4 and H8 (Table 3). The quadrats in which E. vigursii was found in the other groups were classified as NVC: U3; M25a — Erica tetralix sub community; H4 and H8b Danthonia decumbens sub community. DIRECT ORDINATION WITH E. VIGURSII USING DETRENDED CORRESPONDENCE ANALYSIS (DCA) The distribution of the floral species from all three sites formed distinct floristic groups with the exception of Erica cinerea, Hypochaeris radicata and Agrostis curtisii (Fig. 2). There was close correlation between A. curtisii and E. vigursii in terms of how frequent A. curtisii was encountered in each of the micro quadrats at all of the sites. When the sites were individually analysed using DCA there were still strong correlations between A. curtisii and E. vigursii (Fig. 2) although not all the species named were present within each site. In Site 1 there were close correlations of E. vigursii with A. curtisii, Potentilla erecta, and Pilosella officinarum. Species which showed as outliers in this site were Lotus corniculatus, Ulex gallii, Thymus spp. and Erica tetralix. In Site 2 E. vigursii was strongly correlated with A. curtisii and P. erecta and associated with Agrostis capillaris, Viola spp. and bare ground whilst the species that show as outliers were Galium saxatile, Carex flacca and Hypochaeris radicata. ECOLOGY OF EUPHRASIA VIGURSII 351 TABLE 2. SPECIES COMPOSITION OF THE THREE COMMUNITY TYPES DEFINED BY TWO-WAY INDICATOR SPECIES ANALYSIS OF THE QUADRAT DATA FROM THE THREE SITES OF EUPHRASIA VIGURSI IN DEVON AND CORNWALL Species Twinspan group 1 2 3 4 5 Ulex gallii V VI VI I 08 Euphrasia vigursit I Il I Il Thymus spp. V VI IV Erica tetralix II VI IV Carex flacca VI IV Il II Il HAypochaeris radicata IV IV II IV Dactylis glomerata I Il Teucrium scorodonia Il Stachys officinalis Il Serratula tinctoria VI VI Viola spp. V VI II I Erica cinerea VI Il Il II Plantago lanceolata II II oI Ulex europaeus IV I V I Agrostis curtisil Il VI VI VI VI Potentilla erecta VI VI VI VI VI Vaccinium myrtillus Il Pilosella officinarum I Il Danthonia decumbens Il Euphrasia micrantha I Calluna vulgaris IV I Polygala serpyllifolia I II Il Euphrasia anglica II II Il Galium saxatile VI II VI Lotus corniculatus II II II Molinia caerulea VI VI IV Hypericum spp. I I Pteridium aquilinum VI VI Senecio jacobaea Il VI Anthoxanthum odoratum Il Holcus lanatus IV Agrostis capillaris I Ill VI Rubus fruticosus II Bare ground Il IV I II Il Dead material VI IV Key to species frequency in each group: 1 = 1-5%; ll = 6-20%; Ill = 21-40%; 1V = 41-60%; V = 61-80%; V1 = 81-100% 352 L. GRANADOS & S. D. LANE TABLE 3. CORRELATION OF NVC COMMUNITIES WITH INDICATOR SPECIES GROUPS AND STUDY SITES Two way indicator species groups NVC communities 1 H6 Erica vagans— Ulex europaeus heath H8 Calluna vulgaris — Ulex gallii heath H4 Ulex gallii — Agrostis curtisii heath Dy H8 Calluna vulgaris — Ulex gallii heath U3 Agrostis curtisii heath 3 U3 Agrostis curtisii heath H4 Ulex gallii — Agrostis curtisii heath H8 Calluna vulgaris — Ulex gallii heath 4 U3 Agrostis curtisii heath U20 Pteridium aquilinum — Galium saxatile community W25 Pteridium aquilinum — Rubus fruticosus underscrub M25 Molinia caerulea — Potentilla erecta mire 5 U3 Agrostis curtisii heath W25 Pteridium aquilinum — Rubus fruticosus underscrub M25 Molinia caerulea — Potentilla erecta mire NVC communities U3 Agrostis curtisii heath H4 Ulex gallii — Agrostis curtisii heath H8 Calluna vulgaris — Ulex gallii heath Site 2 U3 Agrostis curtisii heath U20 Pteridium aquilinum — Galium saxatile community M25 Molinia caerulea — Potentilla erecta mire W25 Pteridium aquilinum — Rubus fruticosus underscrub Site 3 H4 Ulex gallii — Agrostis curtisii heath H6 Erica vagans — Ulex europaeus heath H8 Calluna vulgaris — Ulex gallii heath U3 Agrostis curtisii heath In Site 3 close associations were apparent between E. vigursii, Hypochaeris radicata and Agrostis curtisii as well as Ulex gallii and Potentilla erecta. Species that were outliers and therefore showed less of an association were Serratula tinctoria, Carex flacca, Erica tetralix and Erica cinerea. ENVIRONMENTAL SITE COMPARISON The macro results for both the UNIANOVA analysis (Potassium and Phosphorus) and for the Kruskal-Wallis test (pH and Nitrogen) indicate significant differences between the sites for environmental variables (P<0Q-05). Differences were observed in pH between the macro and micro-quadrats in Site 2 and Site 3 but not in Site 1 (Fig. 3). However, mean values for the micro quadrats were higher than for the macro quadrats at all three sites. On the sites surveyed, EF. vigursii distribution was strongly linked to soil micro-environment, germination correlating with a pH range from 3-3 to 4-8, a Potassium range of between 15 mg/100 g to 53:60 mg/100 g, a Phosphorus range between 0:04 mg/100 g and 7:44 mg/100 g and a Nitrogen range between 0-34% and 9-62%. Results of the UNIANOVA analysis indicated a significant difference in Phosphorus levels (F = 7-446, P>0-05) but the results for Potassium were not significant. Results of the Kruskal-Wallis test indicated a_ significant difference in pH (P<0-05) and in Nitrogen (P<0-05) between the spatial scales (micro and macro). Chemical comparisons of _ the individual sites at micro and macro levels revealed significant differences in Phosphorus levels at Site 1 (t-test, P<0-05), and pH levels at Site 2 and Site 3 (Kruskal-Wallis test, P<0-05). No significant differences were seen in the other parameters recorded. Comparing these results with the descriptive vegetation analyses suggests that E. vigursii distributions generally link with higher levels of nutrients and pH. ECOLOGY OF EUPHRASIA VIGURSII Viola spp —- Lotus corniculatus + Thymus spp - Agrostis curtisti ++ ee) Nn eS) Erica cinerea + Hypochaeris radicata + Euphrasia vigursii re a + p Poa sp Galium saxatile Pilosella officinarum Agrostis capillaris-e = Potentilla erecta Carex flacca + Erica tetralix 4 Ulex gallii Senecio jacobaea = Serratula tinctoria =F Hypericum spp FIGURE 2. Combined Ordination plot of total floristic species associated with Euphrasia vigursii from Detrended Correspondent Analysis of the ninety micro quadrats in Site 1, Site 2 and Site 3. DISCUSSION FLORISTIC COMMUNITY The phytosociological analysis indicates that E. vigursil 1S a species associated with heath and acidic grassland communities typically containing Ulex gallii, Agrostis curtisii, Calluna_ vulgaris together with Molinia caerulea and Potentilla erecta mire. In all the sites there are external influences which reduce the plant communities’ ability to transpose to successional community processes. Colonisation in these communities by plants is limited to those species that are char- acteristically tolerant of environmental stresses, low nutrient availability and low soil pH (acidity), which in turn is associated with high Aluminium and Iron toxicity (Crawley 1997). Acidification of soil is a result of influences of the base substrate by mineral weathering, temperature, hydrology and leaching (Crawley 1997). Particular plant species can acidify the immediate edaphic environment, for example Calluna and Ulex species produce strong acidifying litter which rapidly reduces the pH of the soil beneath them (Grubb & Suter 1971). ASSOCIATED FLORA OF E. VIGURSII Germination of FE. vigursii within the microenvironment was closely correlated with the presence of A. curtisii as opposed to its other known host species, U. gallii. Studies by Silverside (1999) however, indicate the opposite — i.e. E. vigursii apparently avoids the more open, species-poor areas dominated by A. curtisii. Silverside does mention, however, that E. vigursii 1s frequently reported as an associate. The species that are associated with E. vigursii (Tables | and 3) have attributes, apart from Carex flacca, for counteracting low nutrient availability by the adoption of possessing VA mycorrhizals (Grime 1996). They therefore have the ability to use normally unavailable complex organic nitrogen sources, making them less dependant on mineralisation of organic matter (Aerts 1999). Nutrient-poor ecosystems are associated with low litter production and slow decomposition that leads to low rates of Nitrogen cycling; the ability to 354 L. GRANADOS & S. D. LANE survive in such environments may prevent the invasion of highly competitive species dependant on high Nitrogen availability (Aerts 1999). E. vigursii does not possess symbiotic associations such as mycorrhizals but can, if resources are inadequate, parasitise other species, forming haustoria to obtain water, mineral nutrients and carbohydrates from the host’s root system. (Lammi et al. 1999). ENVIRONMENTAL VARIABLES pH Significant differences in pH were evident between macro and micro environments in sites 2 and 3 with levels ranging between 3-4 and 4-35. In Site 1 pH levels for both macro and micro samples were intermediate between these ranges and not significantly different, which suggests that in Site 1 any micro site would be suitable for germination in terms of pH preference. pH levels can be a good indicator of the nutrient availability to plants in a soil and previous work indicates that soils may show small scale heterogeneity in pH levels over small distances (Crawley 1997). This has been confirmed in this study. Phosphorus In Site 1 E. vigursii had germinated in a microsite significantly higher in Phosphorus than the macro environment (Fig. 3) suggesting a preference for Phosphorus levels that range from 0:26 mg/100 g to 19-01 mg/100 g. Phosphorus and Nitrogen are considered the two limiting factors which control plant growth and are geo-chemically complex (Zheng et al. 1999). There was a significant difference between the sites; in particular Site 1 had generally low levels of Phosphorus. In environments with low pH levels Phosphorus availability can be limited. However, if pH levels are increased Phosphorus utilisation improves more markedly than that of Potassium or Nitrogen (Rorison 1971). Low pH levels are associated with deficiency in Calcium, an element that renders phosphoric acid unavailable to plants and which might account for the lower levels observed. (Crawley 1997). Potassium E. vigursii germinated in a micro environment with Potassium levels ranging from 26:2 mg/100 g to 52:5 mg/100 g. Although differences between the macro and micro environment for each site were not statistically significant, in Site 1 the mean Potassium levels ~° were slightly lower in the micro environment. There was a significant difference between the sites (Fig. 3) and the mean levels were higher in Site 1 than in Site 2 and Site 3, which could be the result of periodic swaling. Higher levels of Potassium have been reported in ash deposits following normal management fires although much is lost through leaching (Legg et al. 1992). Nitrogen No significant differences between Nitrogen supply at macro and microsite level were evident at any of the sites, but there was a definite trend for increased levels where E. vigursii had germinated. There were significantly higher Nitrogen levels in Site 1, possibly linked to the grazing regimes in place. Nitrogen levels in grazed swards fluctuate with patterns of defoliation and severity of grazing | determines the relative importance of the two Nitrogen recycling pathways: internal recycling — remobilization from senescent leaves; and external recycling — via animal intake and urine and faeces return (Lemaire & Chapman 1996). The former process is subject to leaching and gaseous loss, increasing the stock rates within grazing systems and thus leading to lower of Nitrogen utilisation efficiency by vegetation when uptake rates are lower than input rates. (Lemaire & Chapman 1996). Spatial distribution of E. vigursti The position in a habitat of an individual plant is determined by its capability for colonisation, competition and longevity and that is dependent initially on its immediate spatial separation from neighbours (Ross & Harper 1972; Tilman 1994). E. vigursii has to recognise, after the arrival of a seed in a micro site, the location of a potential host species which is an important factor in determining its survival; it does not have a persistent seed bank (Grime 1996). Studies on species of Euphrasia in cultivation indicate that when a seed is planted 2-5 cm from a host, it establishes a week earlier than those spaced 4 5 cm and two weeks earlier than those spaced 6-5 cm apart; the latter also result in lower percentage survival (Yeo 1964). In addition the seeds are very small (<1 mm) and wind dispersed (Stace 1999) and upon landing are subjected to a highly heterogeneous micro soil surface. As a result they may require a particular morphological dispersal site in order to establish (Harper er al. 1965). The soil solution contains dissolved substances that can affect germination, in a) ioe) ECOLOGY OF EUPHRASIA VIGURSII % Phosphate an Se Ce ee % Potassium Nitrogen % FIGURE 3 (a) Soil pH, (b) Phosphorus, (c) Potassium and (d) Nitrogen values in Macro and Micro Quadrats in Site 1, Site 2 and Site 3. Vertical bar = + standard deviation. 356 L. GRANADOS & S. D. LANE particular allelochemicals that inhibit germination; but the host plants of parasitic angiosperms produce chemical signals that stimulate germination of the parasitic seeds and allelochemicals at low concentrations may also be stimulatory (Karssen & Hilhorst 1992). It is possible that nutrient and pH levels may also indicate the spatial position in relation to a host species. The subsurface of the edaphic environment contains a matrix of root systems and although in this study E. vigursii has been found to be closely associated with A. curtisii, 1t remains possible that new young plants of Ulex gallii were also forming but were not detected above ground level. Euphrasia species show a particular preference for Leguminosae over Gramineae because of increased Nitrogen fixation (Yeo 1964). Competition The species associated with EF. vigursii are predominately of low or retarded growth, either because they have an ascending growth form or their vigour is kept in check by grazing or abiotic influences. A _ separate study by Dartmoor National Park (Norman Baldock, Ecologist Dartmoor National Park; unpublished data 1997) targeted a 10 m x 10 m grazing exclusion site close to Site 1. This reported that over a seven year period shrubby species such as Ulex and Calluna formed a dense canopy reducing the frequency and abundance of low herb species. Heathland communities are a direct result of disturbance by natural stochastic events or anthropogenic man- agement regimes and are considered an essential resource for seasonal grazing of livestock by farmers. Disturbance of heath and grassland communities by fire and grazing blocks succession to dominant competitive species (Smith & Rushton 1994). Selective grazing produces a patchy distribution of vegetation and irregular edges, creating gaps (Hearn 1995; Bullock et al. 1994) which can be utilised by species which require high light availability and are poor competitors. This is considered an essential factor in successful colonisation of E. vigursii (Yeo 1964). The main associate with E. vigursii is Agrostis curtisii, which can be found in a number of important heathland communities Where it can form dense patches and is considered unpalatable to herbivores. It is highly competitive with other Poaceae species, not by its ability to compete for light but its ability to reduce possible establishment sites for other Poaceae species due to its dense sward; it is also known to become especially abundant after burning (Rodwell 1992) together with Ulex species (Grime 1996). Euphrasia species and rare species in general and are considered superior competitors to common grasses (Rabinowitz 1981). Potentilla erecta can have either a prostrate or ascending growth form which reduces the damage caused by trampling and is not as readily grazed, affording a competitive advantage in grazed situations (Hearn 1995). Similarly E. vigursii also overcomes possible competitive interactions with other low growing species by forming dense clusters of individual plants (Yeo 1964). Inter and intra- specific interactions between individuals may occur in E. vigursii, as Yeo also reports that — Euphrasia species can parasitise more than one host at a time — for example Carex curta, Poa annua and Trifolium pratense, and also members of its own species. Many endangered endemic species have low population numbers, together with small geographical ranges and narrow habitat speci- ficity which can cause natural selection to favour traits which offset the disadvantages of small local population size (Rabinowitz 1981). However, natural selection can also reinforce narrow habitat selection imposing further distribution. restrictions (Rosenzweig & Lomolino 1997). E. vigursii is able to coexist with species that have adapted to benefit from an environment that is otherwise hostile to many of the common competitive species and its habitat preference and choice is a result of evolutionary habitat selection (Bazaz 1991). 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Cambridge University Press, Cambridge. TILMAN, D. (1994). Competition and biodiversity in spatially structured habitats. Ecology 75: 2-16. YEO, P. F. (1964). The growth of Euphrasia in cultivation. Watsonia 6: 1-24. ZHENG, D. W., AGREN, G. I. & BENHGTSSON, J. (1999). How do soil organisms affect total organic nitrogen storage and substrate nitrogen to carbon ratio in soils? A theoretical analysis. Oikos 86: 430-442. (Accepted July 2006) Watsonia 26: 359-371 (2007) 359 Clonal diversity in British populations of the alien invasive Giant Knotweed, Fallopia sachalinensis (F. Schmidt) Ronse Decraene, in the context of European and Japanese plants C. H. PASHLEY'’ Institute for Lung Health, Department of Infection Immunity and Inflammation, University of Leicester, Glenfield Hospital, Groby Road, Leicester, LE3 9QP J. P. BAILEY’? and C. FERRIS Department of Biological Sciences, University of Leicester University of Leicester, University Road, Leicester, LEI 7RH ABSTRACT Fallopia sachalinensis (F. Schmidt) Ronse Decraene was introduced to Europe during the 19th century, and whilst not as invasive as the notorious F. Japonica var. japonica (Houtt.) Ronse Decraene, it is still an invasive plant capable of both clonal and sexual reproduction. It is also a significant pollen source for the male-sterile F. japonica var. japonica, producing the highly invasive hybrid F. xbohemica (Chrtek & Chrtkova) J. P. Bailey. Given the lack of diversity in F. japonica var. japonica, F. sacha- linensis is a potentially important source for introducing novel variation into F. xbohemica. The majority of British F. sachalinensis was found to be one of two widespread genotypes, either a male- fertile or a male-sterile clone. In contrast there was a much higher level of genetic variation detected in both the rest of the introduced range and in the native range. Evidence is given for two _ separate introductions of native material into Britain; one from Niigata, Honshu, directly into Britain, and the other from Northern Japan, via St. Petersburg (Russia) to Europe. KEYWORDS: genetic variation, introduced plants, Giant knotweed, Polygonaceae. INTRODUCTION The introduction of plants to new environments often brings together species that may not have been sympatric in their native range. These factors can lead to interspecific hybridisation occurring between a native and an invading plant species, or two invading species. This ‘Corresponding author present address e-mail:chp5 @leicester.ac.uk “e-mail: jpb @leicester.ac.uk hybridisation can have numerous effects, the extremes being the evolution of new plant taxa e.g. Senecio cambrensis Rosser, which arose as an allohexaploid hybrid between Groundsel, S. vulgaris L., and the introduced species, Oxford Ragwort, S. squalidus L. (Harris & Ingram 1992), or the extinction of the native species (Sakai et al. 2001). Genetic variation may be lower in newly established introduced populations because either the number of plants introduced is often small or genetic drift during colonization may reduce genetic variation in the newly established population. Thus a newly establish- ed population is likely to be much less genetically diverse than the population from which it is derived (Sakai et al. 2001). When an introduced ornamental plant can be easily propagated by vegetative means, such as is the case with Japanese Knotweed sensu lato (s.1.) the likelihood is that very little material is introduced at any one time. Clonal growth-is a widespread phenomenon in the plant kingdom, which can also occur in lichens, fungi and some groups of lower animals. Plants can reproduce clonally in many ways, including the formation of ramets on creeping stems above or belowground, and the vegetative production of plantlets and bulbils on aerial plants (Stuefer et al. 2001). For the plants that comprise the group Japanese Knot- weed s./., clonal spread is achieved through large underground rhizomes, and in the case of the most renowned member of the group, 360 C. H. PASHLEY, J. P. BAILEY & C. FERRIS TABLE 1. LOCATION DETAILS OF F. SACHALINENSIS, SEX OF THE PLANT WHERE KNOWN, CHLOROPLAST HAPLOTYPE (MPH) AND GENOTYPE (G) RESULTS Plant ID B9 B10 Bll B12 B13 B14 B15 B16 B17 B18 B19 B20 B21 B22 B23 B24 B25 B26 B27 B28 B29 B30 B31 Gl G2 G3 G4 G5 Cl C2 Ul U2 Jl J2 J3 J4 J5 J6 V7 J8 J9 J10 MPH f= (Ni. ON ON, ON (ON (ON “ON ON (ON) 1" 0) ON (ON (ON ON ON (ON ON (ONION (ON (ON (ON ON (ON ON ONION ON On ON G il il il i4 il il i4 i4 il il il 16 16 i4 i4 nlo milul nl5 ns n5 nl2 n3 nl4 Location Caerynwich, Water gardens, Wales Caerynwich, Wales Congerstone, Leicestershire, England Wanlip, Leicestershire, England Amroth Beach, Wales Amroth, Wales Cwrt Newydd, Wales Howey village, Wales Nant Y Frith, Wales Cirencester Roundabout, England Cirencester Abbey grounds, England Guys Cliff, Warwick, England Guys Cliff, Warwick, England Epwell, Warwicks, England Nuneaton, Warwickshire, England Tiscott, Cornwall, England Coed-y-felin Woods, Wales Scarrier, Cornwall, England Polzeath, Cornwall, England Christchurch road, South Hampshire, England Hempstead Old Rectory, England Colchester, Bunting’s nursery, England Colchester, Bunting’s nursery, England Aachen, Germany Aachen, Stadtwald, Germany Aachen, Stadtwald, Germany Leverkusen, Germany Umminger, Germany Konopiste, (Benesov), Czech Republic Kostelec, (Kolin,), Czech Republic Aberdeen, Washington, USA 125th Street, Seattle, USA Joetsu, Niigata, Honshu, Japan Joetsu, Niigata, Honshu, Japan Yonezawa Rd, Fukushima, Honshu, Japan Yonezawa Rd, Fukushima, Honshu, Japan Zao Quasi National Park, Yamagata, Japan Mt. Aoba-yama, Miyagi, Honshu, Japan Yamagata, Japan Eai River Valley, Miyagi, Honshu, Japan Hirosaki, Iwaki river, Aomori, Honshu, Japan Hirosaki, Iwaki river, Aomori, Honshu, Japan Grid reference/ location SH7518 SH7518 SK3605 SK5919 SN1608 SN1608 SN4947 SO00558 S02654 SP0201 SP0202 SP2865 SP2865 SP3540 SP3592 SS2309 ST1882 SW7244 SW9378 SZ2392 TG4028 TMO0025 TMO0025 N 37 06 ES Sab N 37 06 ENS sels IN 3 S03 7/ E 140 22.36 N37 5037 E 140 22.36 NB SiO7A/s E140 26.19 N 38 15.00 E 140 53.14 N 38 38.97 E 140 10.36 N 38 44.05 E 140 46.12 N 40 36.00 E 140 26.37 N 40 36.00 E 140 26.37 Sex male-fertile male-fertile male-fertile male-fertile male-fertile male-sterile male-sterile male-fertile male-fertile male-fertile male-sterile male-sterile male-sterile male-sterile male-sterile male-sterile male-fertile male-fertile male-sterile male-fertile male-fertile male-fertile male-sterile male-sterile male-sterile male-sterile male-fertile male-fertile male-fertile male-fertile male-sterile male-fertile CLONAL DIVERSITY IN BRITISH FALLOPIA SACHALINENSIS 361 TABLE 1 CONTINUED Plant ID MPH G___ Location Grid reference/ Sex location Ut 6 nl Memuro, Hokkaido, Japan N 42 50.12 E 143 00.44 , 6 n7 Memuro, Hokkaido, Japan N 42 50.12 E 143 00.44 w13 6 n6 Memuro, Hokkaido, Japan N 42 50.12 E 143 00.44 J1i4 6 n4_Ishikari, Hokkaido, Japan N 43 12.24 E 141 23.11 Tiles 6 n9_ Ishikari, Hokkaido, Japan N 43 12.24 1B, WA BI J16 6 nl6 Ishikari, Hokkaido, Japan N 43 12.24 Je, NAN) Z3)all i — genotype possessed by a plant from the introduced range n — genotype possessed by a plant from the native range Fallopia japonica var. japonica (Houtt.) Ronse Decraene, as little as 0:7g is required to give rise to a new plant (Brock & Wade 1992). Well-established plants develop woody stocks with a central taproot that penetrates vertically into the ground. Buds form on the stock and woody rhizomes between autumn and winter, and emerge to give vertical shoots the follow- ing spring. Woody stocks continue to increase in width by secondary thickening with age, and also produce lateral creeping rhizomes within their first year (Beerling et al. 1994). The rhizomes of a mature plant can extend up to 7m away from the parent plant (Child & Wade 2000). Japanese Knotweed s./. comprises taxa from the genus Fallopia Adans., section Reynoutria (Houtt.) Ronse Decr. This includes F. japonica, which in Britain can be found as both var. Japonica and var. compacta (Houtt.) Ronse Decr. (Hook. f.) J. P. Bailey; F. sachalinensis (F. Schmidt) Ronse Decr., which is also commonly known as Giant Knotweed; the hybrid between F. japonica and F. sacha- linensis called F. xbohemica (Chrtek & Chrtkova) J. P. Bailey; and any backcrosses these plants may form (Bailey & Conolly 2000). The different species were introduced to Britain from parts of Asia at various times during the 19th century whilst the hybrids are believed to have arisen since the introduction of the parental species. Fallopia japonica var. japonica, the most common of the Japanese Knotweed s./. plants in Europe, was shown to be represented in Britain by a single male-sterile octoploid (2n = 88) clone. This clone was also found in France, Germany, the Czech Republic and the U.S.A. (Hollingsworth & Bailey 2000a), and in Holland (Pashley 2003). In contrast there appeared to be higher levels of genetic variation within the UK in the related Giant Knotweed, F. sachalinensis (Hollingsworth 1998; 2000b), than in the other introduced Fallopia species. Fallopia sachalinensis was first recorded in the wild in Germany in 1869, in the Czech republic in 1869, and in Great Britain in 1896 (Sukopp & Starfinger 1995). Examination of historical plant collection and distribution records suggest that there were at least two routes by which F. sachalinensis may have arrived in Europe. The first is via St. Peters- burg, Russia, and the second route is via Kew Botanic Gardens, London (Bailey & Conolly 2000). F. sachalinensis has not spread to the same extent as F. japonica var. japonica. Instead, its occurrence appears to reflect many independent primary escapes (Conolly 1977). F. sachalinensis may not be as invasive as F. Japonica var. japonica, but it can be a signi- ficant pollen source for the male-sterile F. Japonica Var. Japonica, producing the hybrid F. xbohemica (Chrtek & Chrtkova) J. P. Bailey. F. xbohemica is gaining in notoriety, and is ~ thought to be at least as invasive as F. japonica var. japonica (Brabec & PySek 2000; Bimova et al. 2001). Given the lack of diversity in F. Japonica var. japonica, and the rarity of F. Japonica var. compacta (Hook. f.) J. P. Bailey 362 © Hs PASHLEY? J. PeBAIEBNG So CSEERRIS (the alternative F. japonica parent for F. xbohemica), the genetic composition of F. sachalinensis is an important factor for potentially introducing novel variation into F. xbohemica. AIMS The main aims of this study were to estimate levels of genetic variation among British F. sachalinensis, in order to determine whether sexual reproduction or clonal spread best explain the current distribution. We have compared genetic diversity within the British plants to a limited number of plants from other countries where F. sachalinensis has been introduced, namely Germany, the Czech Republic, and the USA, to see if the genetic diversity within Britain is similar or different from the rest of the introduced range. We have investigated the levels of genetic variation among native F. sachalinensis from Japan in order to compare levels of genetic diversity between the introduced and native range, and to potentially determine the region of Japan from which the introduced plants originated. MATERIALS AND METHODS PLANT MATERIAL Leaf material was obtained from 56 F. sachalinensis accessions, 40 from _ the introduced range and 16 from the native, as detailed in Table 1. The introduced plants include 31 individuals representing 21 populations from Britain, five from Germany, two from the Czech Republic and two from the USA. The 16 native plants were chosen to represent the geographical distribution of the plants within Japan. Voucher specimens for each accession were collected and deposited at the University of Leicester herbarium (LTR). DNA EXTRACTION Total genomic DNA _ was isolated from approximately 100 mg of fresh leaf tissue using the DNeasy plant mini kit (QIAGEN, Valencia, CA). Duplicate extractions were prepared for each plant. CHLOROPLAST RFLP ANALYSIS The chloroplast regions trmC-trnD and trnF- trnV were amplified using the universal primers of Demesure et al. (1995) and Dumolin-Lapegue ef al. (1997) respectively. PCR reactions were carried out in a total volume of 25 uL containing 2:5 ul 10 x NH, reaction buffer (TaKaRa), 2-5 ul dNTPs (2 mM), 0:5 ul of each primer (10 uM), 0-5 units of Ex taq polymerase (TaKaRa) and 2:5 ul of genomic DNA (1 ng/ul). Amplification was carried out using the following PCR profile: 1 cycle of 94 °C for 4 min; 32 cycles of 45s at 94 °C, 45s at 58 °C, and 2 min 30s¥at) 72a Gea soak at 72 °C for 10 min; and finally held at 10 °C for 30 min. Chloroplast PCR products were digested for a minimum of one hour at 37 °C, with the restriction enzyme Hinfl following the method of Ferris et al. (1993). 10 ul of the restriction digest products were then run on 3% MetaPhor XR agarose gel for trnC-trnD products or 1-6% multipurpose agarose gel for tmF-trnV. Fragments were visualized by staining with ethidium bromide (0:5 ug/ml). INTER-SIMPLE-SEQUENCE REPEAT (ISSR) ANALYSIS Thirty primers from the University of British Columbia (UBC) primer set nine and twelve from the laboratories of Professor Mike Wilkinson (University of Reading) were screened. A final set of seven polymorphic primers was selected. on the basis of reproducibility between duplicate DNA extractions from the same plant, and between different PCR runs (details available on request). PCR reactions were carried out in a total volume of 25 uL containing 2:5 ul 10 x NH, reaction buffer (Bioline), 2°5 ul dNTPs (2 mM), 1:25 ul magnesium chloride (50 mM), 0-33 ul of primer (15 UM), | unit of BIOTAQ DNA polymerase (Bioline) and 10 wl of genomic DNA (1 ng/ul). Amplification was carried out using the following PCR profile: 1 cycle of 94 °C for 4 min; 40 cycles of 20s at 94 °C, 30s at X °C, and 1 min at 72 °C; a soak at 72 °C for 7 min; and finally held at 4 °C for 30 min. X is primer specific (details available on request). Reactions were performed in duplicate using the independently extracted DNA, and 10 ul from each reaction, with duplicates placed in adjacent wells, were run on a 1-6% agarose gel with a | kb ladder in the outer wells. Products were visualized by staining with ethidium bromide (0:5 Ug/ml). Bands were considered to be reproducible when the same DNA pattern was obtained using the two different DNA isolates. A table of presence/absence of ISSR fragments was constructed and is available from _ the corresponding author on request. CLONAL DIVERSITY IN BRITISH FALLOPIA SACHALINENSIS 363 DATA ANALYSIS RFLP bands were scored by eye and recorded in a binary data matrix table, as presence (1) and absence (0). Each combination of these bands for a given chloroplast product is referred to as a haplotype. The combination of these haplotypes is referred to as a multi- primer-haplotype (MPH) and is intended to represent the chloroplast type of the plants in which it was found. Winboot (Yap & Nelson 1996) was used to bootstrap the ISSR binary data 1000 times using the Jaccard’s coefficient to produce confidence values. The companion program WinDist was then used to produce a matrix of pair-wise genetic distances between all individuals using the complement of the Jaccard’s similarity coefficient. A Neighbour- Joining tree was drawn from the resulting matrix using the NEIGHBOUR option in PHYLIP version 3.6a3 (Felsenstein 2002) and TreeView version 1.6.6 (Page 1996). Addition- ally, the number of differences between each ISSR genotype was calculated and analysed using MINSPNET (Excoffier & Smouse 1994) to produce a minimum spanning tree. RESULTS CHLOROPLAST HAPLOTYPES Six different MPH were detected from the native plants, designated MPH 1-6. All 47 of the introduced F. sachalinensis accessions were found to have a MPH also found in the native samples, 44 having MPH 6 and the remaining three having MPH 1 (Table 1). ISSR GENOTYPES Each ISSR primer generated a banding pattern referred to as a phenotype. From five to thirteen bands were scored per _ primer, resulting in between five and twenty-eight phenotypes depending upon the primer. A total of fifty-nine bands was scored, of which 52 were polymorphic. A unique combination of these phenotypes is assumed to be representative of a genotype. A total of 32 genotypes were detected (Table 1). Sixteen genotypes were found in plants from the introduced range; these have been given the letter “1” followed by a unique number. The remaining sixteen genotypes were from native Japanese plants; these have been given the letter “n’” followed by a number. No native genotypes were found in the introduced plants. DISTRIBUTION OF GENOTYPES Sixteen native F. sachalinensis plants were analysed. As seen in Fig. 1, each plant possessed a unique genotype, indicating a high level of genetic diversity among native plants. Plants from the same geographical area share the same chloroplast haplotype, however none from the same area shared a genotype. Whilst some plants within populations were genetically similar (Fig. 2, MPHs 1, 3, and 4), this was not true for all of them. By contrast, in Britain only eight genotypes were found from the thirty-one plants analysed (Fig. 3). As can be seen there were two widespread clones, il shared by eleven accessions and 14 by thirteen. These represent a male-fertile and a male-sterile clone respect- ively, both with chloroplast MPH 6. The remainder of the genotypes were found at single localities, with a maximum of two plants sharing the genotype. The three accessions with MPH 1 were found to contain different ISSR genotypes. The other three genotypes with MPH 6 were represented by a male-fertile plant from South Hampshire (113), two male- sterile stands from Warwick (i6), and a single stand from Merioneth (12). There were nine F. sachalinensis accessions analysed from the introduced range that were not from Britain. The two plants from the U.S.A had unique genotypes (i7 and 112). Five plants came from Germany, three of which were from the same area of Germany, Aachen, however all five were found to have unique genotypes (13, 19, 110, 114 and i115). Two plants were analysed from the Czech Republic. Whilst one had a unique genotype, 15, the other possessed the male-sterile genotype, 14, which was common in Britain. NEIGHBOUR JOINING TREE The Neighbour Joining tree for all genotypes is shown in Fig. 4. The internal branches are short and the majority of the distances are confined to the terminal branches. Bootstrapping 1,000 times has shown that only four of the branches were supported over 50% of the time. In each case the support was for a group of two genotypes. There was support of 68:0% for the two genotypes with MPH 4, native plants from Fukushima (nl0O and nll), and a bootstrap value of 91-5% for the two native genotypes with MPH 1, which came from Niigata (n2 and nl3). The other support was for introduced plants. The genotypes i8 and ill (76:9% bootstrap) consisted of two of the three British 364 C.H. PASHLEY, J. P- BAILEY & CoFERRIS 4 Lo ees Honshu MPH 1 MPH 2 MPH 3 MPH 4 MPH 5 MPH 6 introduced genotype native genotype FIGURE 1. Localities of Japanese F. sachalinensis genotypes. The map was produced using Dr. A. J. Morton’s DMAP program. A simplified version of the unrooted Neighbour-Joining tree (Fig. 4) has been included to show the relationship between the different native genotypes, and their relationship to the introduced plants. 365 CLONAL DIVERSITY IN BRITISH FALLOPIA SACHALINENSIS UMOUY dIOYM poeoipul st odAjouss ay) Jo xag ‘adAjouas oy} YIM spuodsa1i09 yey) adAjojdey jsepdosopyo oy) syuososdar adAjouas oy) JOA posodurtiadns adeys oyy, ‘sodAjoues usamjoq soarjeusaye Ajoyty Ay[enbo ose syury Aols oy, “SQOUDIOJJIP JO Joquinu sy} MOYsS syUT]-ssoso sy TZ, ‘sodAjouss sisuauIYYIVS “J UIIMJOQ SddUdIOJJIP JO JaquINU oY) SUIMOYS YJOMJOU SuTUUeds WNUTTUTPY °7 AYNOL i ese ~@) a 1h Benen 9 ey = aH 49}-9e A aqL4955-9T 8A] adAjouas paonponui P adAjouas aaueu g ui g udu OC cydu 22 pydu 5 ¢ ydu ZT ydu | udu oS (ru) 366 C. H. PASHLEY, J. P. BAILEY & C. FERRIS C Crech Republic ‘ U GG G Germany \ eee N Native L_| MPH 1 U USA C) MPH6 i introduced genotype genotype i4 (male-sterile) genotype il (male-fertile) FIGURE 3. Localities of British F. sachalinensis genotypes. The map was produced using Dr. A. J. Morton’s DMAP program. The position of some genotypes have been moved slightly, to allow all samples to be seen, due to multiple records in single recording 10 km squares. A simplified version of the unrooted Neighbour- Joining tree (Fig. 4) has been included to show the relationship between the British genotypes, and their relationship to the non-British introduced plants, and the native plants. CLONAL DIVERSITY IN BRITISH FALLOPIA SACHALINENSIS 367 i10 [| MPH 1 \. MPH 2 “> MPH 3 Cc 2 MPH4 ~~ MPH 5 (© MPH6 introduced genotype native genotype FIGURE 4. Unrooted Neighbour-Joining tree depicting relationships between the various ISSR genotypes, based on Jaccard’s similarity coefficient. Numbers along branches are confidence values based on 1,000 bootstraps. Only bootstrap values greater than 50% are shown. The shape superimposed over the genotype represents the corresponding chloroplast haplotype. plants with MPH 1: the male-sterile plant from Colchester and a plant of unknown sex from Glasgow. The other introduced genotypes that grouped together were il and i2 (99:2% bootstrap) representing the common male- fertile genotype (il) and an individual stand from Merioneth. The majority of the introduced genotypes, twelve of the 16, were all found to diverge from a single branch with no native genotypes forming part of the group, although this group is not statistically supported (Fig. 4). In contrast the native genotypes appeared to arise from four main branches, with two of the branches closer to the branch with the majority of the introduced genotypes than the other two. One of the four branches was further divided into two clear groups, one comprising n2 and nl3, and the other made up of nl, n6, n7 and n8. The three introduced genotypes from plants with MPH 1 were found intermingled with native genotypes, two of them grouping with the Japanese plants with MPH | as well as a genotype, 19, that came from a plant with MPH 6. The other genotype that was found mixed in with the native genotypes was the British male- fertile genotype with MPH 1, 116. MINIMUM SPANNING NETWORK The minimum-spanning network (Fig. 2) is based on the number of band differences between each pair of genotypes. Next to the genotype is a symbol to represent whether the genotype is male-fertile or male-sterile. There was no obvious separation between male-fertile and male-sterile genotypes. The network is almost linear in shape, with only seven equally likely alternative links throughout the whole network. The genotypes found in_ the introduced plants occupy either end of the 368 C. H. PASHLEY, J. P. BAILEY & C. FERRIS network, with six native genotypes found between the nearest of the introduced genotypes at either end. These six genotypes represent 65 band differences between i113 and il6, which were the closest of the genotypes from the two ends. The two common British genotypes (il and 14) were closely related and were found to have only ten band differences. The group of twelve introduced genotypes to the left of the network were the same genotypes that came from the single branch in the Neighbour Joining analysis (Fig. 4). The minimum-spanning network showed a closer relationship between the four introduced genotypes that were not grouped with the other introduced genotypes than did the Neighbour Joining analysis, being interconnected by genotype 111. DISCUSSION DISTRIBUTION AND GENETIC DIVERSITY OF THE NATIVE PLANTS Sixteen native F. sachalinensis plants were analysed, each of which possessed a unique genotype. The native plants harboured a much higher level of genetic diversity than the introduced plants, a common phenomenon found in many different species as a result of the genetic bottleneck experienced during the introduction process (Barrett & Richardson 1986; Barrett & Shore 1989). Whilst plants from the same geographical area shared the same chloroplast haplotype, there was less geographic structuring evident with the genotypes, although there does appear to be some correlation between genetic relationship and geographical distribution. Japanese individuals with MPH 1 came from Niigata, Honshu, and had genotypes n2 and n13, which form a supported group (91:5% bootstrap) on the Neighbour-Joining tree (Fig. 4), and are each others nearest neighbours on_ the minimum-spanning network (Fig. 2). Likewise the individuals with MPH 3 from Aomori, Honshu cluster together as do those with MPH 4 (68% bootstrap) from Fukushima, Honshu. In general, the taxa to the left of the minimum- spanning network as it is portrayed in Fig. 2 were found in the northernmost part of Honshu and on Hokkaido, and those to the right in the central and southernmost part of the native range (Fig. 1). DISTRIBUTION AND GENETIC DIVERSITY OF THE INTRODUCED PLANTS The majority of the introduced genotypes, twelve of the 16, were found to group together both on the Neighbour-Joining tree (Fig. 4) and the minimum-spanning network (Fig. 2). The four introduced genotypes not clustering with these being the three British genotypes with MPH 1 and a male-sterile German plant (G3) with genotype 19 and MPH 6. It is interesting to note that, on both the Neighbour-Joining tree (Fig. 4) and the minimum-spanning network (Fig. 2), the non- British introduced genotypes appear to be more closely related to each other than they do the other genotypes found in Britain (genotypes 13, i5, 17, 110, 112, 114 and 115), the exception being genotype 19. Genotype i13, the British genotype most closely associated with the non- . British introduced genotypes in both analyses being found in Britain in a plant from the closest collection point to mainland Europe. The lack of support for this observation could, however, imply that this is mere coincidence. The odd non-British (German) introduced genotype, 19, having the same chloroplast haplotype (MPH 6) as the majority of the introduced F. sachalinensis, yet has a genotype more closely related to the British introduced plants with the rarer MPH 1. The reason for this is unclear and further sampling and analysis would be required to better understand how this individual arose. The sharing of the male-sterile genotype between Britain and the Czech Republic implies that the majority of the plants (those with MPH 6) were introduced from the same collection, presumably from St. Petersburg botanic garden. There also appears to be a higher level of genetic variation within the non-British introduced F. sachalinensis individuals, as demonstrated by each having a unique genotype, although further sampling of mainland Europe and USA would be required to support or refute this point. Spread of two clones, a male-fertile and a male-sterile clone, appears to be the primary explanation for the current F. sachalinensis distribution within Britain (Fig. 3). The male- sterile clone, i4, being found as far north as Fort William, Scotland, and as far south as Cornwall, England. The male-fertile clone, il, appears to have a more restricted distribution, being found predominantly in the centre of England, and Wales, but spreading from the CLONAL DIVERSITY IN BRITISH FALLOPIA SACHALINENSIS 369 East to the west coast. The remainder of the British genotypes were found at single localities, with a maximum of two _ plants sharing the genotype. Unlike in Britain, the remainder of the introduced plants demonstrate no signs of clonal spread, although as noted above they do appear to be closely related to each other. This greater diversity could be the result of in situ sexual reproduction, but further sampling would be needed to support or refute such a claim. A largely unpublished preliminary study cited by Hollingsworth & Bailey (2000b) also found more clonal diversity in F. sachalinensis (14 genotypes from 30 samples) than in the uniclonal F. japonica var. japonica. However, unlike the current study no shared genotypes were detected between populations. It is probable that in the previous study the lack of shared genotypes among at least some of the sites 1s erroneous and is attributable to PCR artefacts (a lack of time prevented duplicate DNA extractions being used to test the reliability of different genotypes; Hollingsworth 1998). SOURCE OF THE INTRODUCED FALLOPIA SACHALINENSIS Both of the MPHs found in introduced material were found in native plants. The most common in the introduced plants, MPH 6, was found only in individuals from Hokkaido. The rarer introduced chloroplast haplotype found in the introduced range only in Britain, MPH 1, was only found in individuals from Niigata, Honshu. The low level of resolution and support found with the neighbour joining analysis means conclusive statements about the relationships between the native Japanese and introduced genotypes of F. sachalinensis cannot be made. Even so, some observations can be made, and tentative suggestions towards how the current distribution arose can be proposed. Whether the genotypes found in _ the introduced F. sachalinensis were present in the native range before the plants were collected, or have arisen by sexual reproduction since their introduction is unclear. It is likely that the situation has arisen from a combination of the two factors. A limited number of genotypes were probably introduced from Japan, most likely Hokkaido or the Sakhalin Island, into St. Petersburg. The most plausible explanation is that a minimum of a male-fertile and a male- sterile clone were sent to Britain from St. Petersburg as established plants rather than seed. These were then distributed throughout Britain to various nursery gardens, and from there to many stately homes and manors, eventually leading to the escapes reflected in the distribution noted by Conolly (1977). Given how easy F. sachalinensis is to propagate from the rhizome, there would be little point in the gardeners germinating and growing seed for distribution. At a similar time, or earlier, plants arising from the same original collection were probably being distributed to Germany, the Czech Republic and other European countries. Fallopia sachalinensis was recommended, especially on the continent, as a forage plant for cattle, as well as being introduced as a riverbank stabiliser (Bailey & Conolly 2000). For stabilisation of riverbanks, mature plants were most likely used, which would pres- umably originate from vegetative reproduction. In addition to its use as a forage plant, F. sachalinensis was also used on large estates for scenic plantings and cover for shoots (Bailey & Conolly 2000). As with riverbank stabilising, the plants introduced for scenic plantings were probably introduced as mature plants. However, seed may have been used to produce the plants for forage, which could explain why there was found to be higher levels of genetic variation on the continent, but these plants all cluster together away from the native, possibly indicating that they were produced from the genetically limited original introductions by sexual reproduction. Presumably material either directly from St. Petersburg, or from elsewhere on mainland Europe was sent to the USA. Three different collections are presumed to have been housed at St. Petersburg: that of Dr H. Weyrich, from Sakhalin in September 1853; that of P. von Glehn in 1861, also from Sakhalin that arrived in St. Petersburg in early 1863; and material of Japanese origin collected by C. J. Maximovicz during 1859-1864 (Bailey & Conolly 2000). Whilst it is not possible, from this study, to determine whether the introduced plants came from Sakhalin or Japan due to a lack of material from the - Sakhalin island, data would suggest that if the plants with MPH 6 came from Japan it would have been from the northern island Hokkaido. The three genotypes found in Britain in plants that have MPH 1 were probably introduced directly from Niigata on Honshu, 370 C. H. PASHLEY, J. P. BAILEY & C. FERRIS Japan. Whilst it is possible that this chloroplast type could be found in non-sampled Japanese sites, a study of chloroplast variation in two hundred and forty Japanese Knotweed s./. accessions from eighty seven sites spanning the four main islands that make up Japan, found this chloroplast type to be restricted to Niigata (Pashley 2003). This could explain why no plants with this chloroplast type have been found anywhere besides Niigata and Britain. The two sites in Britain where these were found were Glasgow and Colchester. The plants (both male-fertile and male-sterile) in Colchester shared MPH 1 and were found growing on the site of an old nursery garden reported to have sold F. sachalinensis (Bailey & Conolly 2000). The site in Glasgow where the individual with MPH 1 (B3) was found is close to a botanic garden, and the implication is that the plants at Colchester and the individual B3 from Glasgow came from a common source. CONCLUSIONS The majority of British F. sachalinensis were found to be one of two widespread genotypes, either a male-fertile or a male-sterile clone. The remaining British genotypes that came from plants that share the chloroplast MPH 6 were presumed to have arisen from the same introduction, as were most of the plants in the other introduced countries included in the study, most likely introduced via _ St. Petersburg. Only one genotype was found shared between Britain and any of the other countries to which F. sachalinensis was introduced. That genotype was the widespread male-sterile clone, and was found in a single accession from the Czech Republic. The three genotypes found in Britain in plants that have chloroplast MPH 1 were probably introduced straight from Niigata on Honshu, in Japan, and were not found in any other country included in this study to which F. sachalinensis was introduced. A high level of genetic diversity was found among native F. sachalinensis, with each of the sixteen plants analysed having a _ unique genotype. There was some correlation between genetic relationship and the geographical distribution of these native plants. There is clearly a higher level of genetic diversity in F. sachalinensis than there is in F. Japonica var. japonica (Hollingsworth & Bailey 2000a), however, there is_ strong evidence that the current British distribution of F. sachalinensis has resulted mainly from the spread of two clones rather than via sexual reproduction. ACKNOWLEDGMENTS We are very grateful to MAARA for the financial support to CHP, and to the British Ecological Society (BES), Botanical Society of the British Isles (BSBI), Botanical Research Fund, and Leicester University who provided funding to enable the collection trip to Japan. We also thank Mark Chapman who provided helpful comments on an earlier version of the manuscript. Our thanks also to Ann Conolly, for her support, encouragement, helpful conversations, and company on field trips. We are also grateful to the many people who aided in the plant collection in Britain, including Tony Leech, Jerry Heath, Peter Jepson, R. Hemming, Caroline Wilson, K. Pyne, James Partridge, G. Hutchinson, Peter Zika, James MacFarlene, Dick Shaw, Harry Evans. Finally we are particularly grateful to our Japanese friends and collaboraters Kazuhito Matsuo, Jun Suzuka, Hiroyuki Shibaike, Takashi Fujita, Teruo Sano and Tatuyoshi Morita, without whose generous assistance in the planning and execution of our visit to Japan, this work would have been impossible to achieve. REFERENCES BAILEY, J. P. & CONOLLY, A. P. (2000). Prize-winners to pariahs—A history of Japanese Knotweed s./. (Polygonaceae) in the British Isles. Watsonia 23: 93-110. BARRETT, S. C. H. & RICHARDSON, B. J. (1986). Genetic attributes of invading species, in R. H. GROVES & J. J. BURDON eds. 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(Accepted June 2006) | ; ; io ee : | i re { jews a | i ' ; r | i a i} : : r | i | ' \) J ‘ | | a i i} 1 . ‘4 i | 4 ' mn wei : | = | | : 7) i i | } J ri La Ww : co if i ¥ 4h y Toke i tlie t ; Pn tae | ; r i i a i 1 iH! | i fall ‘ | i i} i ; Ff nal . : } ; ‘atin 1 i} on | i ial { : Soa || ni =) | | - ; alin i ' ¥ i i] i , ; ‘ : we he He} ay h | 1 | _ ; | 1 ; yy fi i i Le? 4 Wy P ' ; f Z ow i] ¥ 1 ! 1 Whi i ; | , | i} 7? J \ | ae i} i . iH} | | i | \ | ‘ | : | i i be | Wi ; i ‘ | ; | \\1)) : 4 { = a | | i f | } Ny hel Hh | r | z He f iM | if aad | Tih N 1 \ i] Z ai 1 | Q Nib} 1H) i i a | | | HAI a | Py , {|| Hi i 4 ete q 4 Oey } ; d vi - u ” Salt I j m ne ti id ; j r ' 1 iY 7 + ; Me 4 a \ i! ‘ -_ 1 H tay 1 iy \ : i 5 ae Tar uy i 1 i i ; ee ; Watsonia 26: 373-379 (2007) Introgression between Plantago major L. subspecies major and subspecies intermedia (Gilib.) Lange. in a British population R. EL-BAKATOUSHI Biology Department, Faculty of Education, University of Alexandria, Egypt A. J. RICHARDS and K. WOLFF School of Biology, Ridley Building, University of Newcastle NE] 7RU ABSTRACT We measured morphological characters of Plantago major from a British population in which subsp. major and subsp. intermedia both occur, and from ‘pure’ populations from different parts of Europe as standards, grown in uniform conditions. In the British population, most individuals fall within the variation of subsp. major, but individuals resembling ssp. intermedia occur as a minority. We attempted to quantify interspecific gene flow between major and intermedia, and tested the hypotheses that the amount of variation can be explained by minority effects, as functions of the relative frequency or density of the two taxa. The first hypothesis is upheld, but evidence for the effect of density is equivocal. A higher rate of gene flow occurs between these two taxa than expected if they are mostly self- fertilised. We suggest that the taxa do not merit specific rank as no breeding barrier seems to separate the species when they become sympatric. KEYWORDS: genetic variation, hybridization, introgression, Plantago, Greater Plantain, plant Speciation, population structure. INTRODUCTION Introgressive hybridization has been defined as the repeated backcrossing of a natural hybrid to one or both parental populations and usually occurs in the direction of one parent only (Anderson 1949). It results in the transfer of genes from one parent to another across a partial breeding barrier (Anderson & Hubricht 1938; Anderson 1949, 1953; Harrison 1993). This can result in one of the parents becoming more variable, having incorporated some genes from the other parent, which may allow the introgressed parent to colonise a new environ- ment. There are some good examples which clearly describe introgression in the literature, where molecular techniques were used to confirm previously doubtful examples. Abbott ef al. (1992) used isozymes to confirm that ray- florets in groundsel Senecio vulgaris had arisen as a result of introgression from the ray- floreted Senecio squalidus. Introgression essentially occurs at the chromosome level. Eshed et al. (1992) revealed that when chromo- somal segments were recombined into the genome of Lycopersicon esculentum (tomato) after hybridization with L. pennellii, the new chromosomal inserts survived through repeated recombination and backcrossing although the size of inserts reduced between the first and the sixth backcross generation. In Plantago, maternally inherited chloroplast DNA (cpDNA) was analyzed by comparing DNA fragment patterns produced by seven restriction endonucleases in four species of Plantago (Hooglander et al. 1993). In this study it was shown that the larger 70bp DNA fragment present in Netherlands Plantago major L. subsp. major (hereafter referred to as ‘major’) but absent in subsp. intermedia (Gilib.) Lange (hereafter referred to as ‘intermedia’), was not observed in a few individuals identified morphologically as major from sites where two taxa grow together. Outside the UK, intermedia and major tend to be separated ecologically, the former taxon being more typical of open saline sites, and has been said to differ biologically, being more often annual or short-lived and self-fertilising than major (van Dijk 1989; van Dik er al. 1988). Thus, it is likely that these taxa are usually isolated spatially in countries such as Denmark and the Netherlands, and more often maintain their integrity there. Nevertheless, Hooglander et al. (1993) suggested that natural successive hybridization and backcrossing onto intermedia mothers had occurred, with the more outcrossing major acting as pollen donors, resulting in intro- gression from intermedia to major. 374 R. EL-BAKATOUSHI, A. J. RICHARDS AND K. WOLFF In a population from north-east England (Metrocentre, Gateshead), morphological clustering in the two taxa major and intermedia is also not clear cut. The aim of the experiments reported in this paper was to investigate the possible occurrence of hybridization and introgression in these mixed populations. We reasoned that the offspring of mothers, some of the seeds of which had been fertilized by pollen from the other taxon or hybrids, would produce offspring that were more variable than those mothers which had not hybridized. By comparing the variability of offspring grown in a standard environment between mothers from different taxa, from areas with differing frequencies of the taxa, and from areas with different densities of adult plants, we were able to investigate parameters which affected rates of hybridisation and introgression. MATERIALS AND METHODS MORPHOLOGY i Four separate field populations (45 plants) were collected near the Metrocentre, Gateshead, UK (NZ220630) and taken into cultivation. The four populations were separated by short distances. Population (A) was collected near the River Tyne which is tidal here. The distance between population (A) and population (B) is 35 m; there is 15 m between population (B) and (C), whilst population (C) and (D) are nearly 40 m apart. Habitats in this area are highly ruderal and are dominated by human activities. Each of the four populations appeared to contain some individuals showing characteristics of both major and intermedia. Although these taxa are reported to differ in up to 12 characters (Hegi 1914, Van Dyk 1984, 1989), average seed number per capsule is the most reliable discriminant, being non-overlapping between the taxa and easily quantifiable (Van Dik 1984, 1989, Morgan-Richards & Wolff 1999). Individuals were classified primarily on the basis of average seed number per capsule, using a discontinuity in the data distribution as the discriminant (El-Bakatoushi 2004) (major < 10-4, intermedia =10:5). This character, which has also been considered an _ important discriminant in all earlier studies of these taxa, is strongly correlated with both major principal components of the variation, and with 7 of the 12 other characters employed (Table 2). Pure populations used for comparison were collected as seed by K. Wolff from districts where only one taxon occurs. These comprised two intermedia populations from Denmark (NYA) and the Netherlands (NPZH) and two major populations from Italy (Salt.2) and the Netherlands (NPZ) (details in El-Bakatoushi, 2004). Using the same cultivated material, El- Bakatoushi (2004) compared characters known to vary between the taxa before and after cultivation, and also examined _ cross- correlation between these characters. As a result of this study, 13 morphological characters were selected which did not change markedly after cultivation and all of which showed some _ significant character state correlation between individuals (Table 2). These characters were scored after two years © growth in standard conditions in pots under glass at Moorbank Gardens (University of Newcastle) (Table 1). Principal Component Analysis (PCA) was used for the multivariate analysis of the morphological characters, using Minitab V.12.1 (1998). PCA is the most mathematically natural ordination technique (Gauch 1982; Digby & Kempton 1987). The use of PCA enabled us to examine multivariate taxonomic relationships in a manner unbiased by choice of character. It was also possible to combine both continuous and multistate characters in the analysis. By means of _ cross-correlation between character states and _ principal components, we were able to examine the relative contribution of ‘each yormihemms characters to these components (Table 2). EXPERIMENTAL HYBRIDIZATION AND HYBRID FERTILITY Plantago major is protogynous, and flowering acropetal so that stigmas are exserted from flowers 1-3 days before pollen is shed (EI- Bakatoushi 2004). Experimental cross- pollination was achieved by applying dehiscing anthers to stigmas exserted from the lowest flowers on a solitary spike, and then excising the remaining flowers and enclosing the cross- pollinated flowers in a_ polythene bag. Reciprocal crosses were made between the two taxa using both individuals from _ the Metrocentre, and the ‘pure’ control pop- ulations. Pollen stainability of samples of 300 grains was examined at x400 magnification using 45% acetocarmine in experimental hybrids and in mother-daughter lines. INTROGRESSION BETWEEN PLANTAGO MAJOR SUBSPECIES Bie) TABLE 1. MORPHOLOGICAL CHARACTERS AND CHARACTER STATES RECORDED FOR PLANTAGO MAJOR AND SUBSP. INTERMEDIA AFTER TWO YEARS CULTIVATION BWNre . Leaf margin 0. Teeth well marked (4+) 1. Teeth slightly apparent (1-4) 2. Undulate toothed 3. Entire . Leaf blade length / length of leaf blade to base from point of maximum width) . Degree of angle at blade / petiole interface (O—180°) . Number of main veins in longest leaf (lower surface of leaf) 5. Leaf indumentum (upper surface, between middle and next vein) 0. Densely covered with hairs 1. Somewhat covered with hairs 2. Slightly covered with hairs 3. Glabrous 6. Leaf indumentum (Lower surface, between middle and next vein) 0. Densely covered with hairs 1. Somewhat covered with hairs 2. Slightly covered with hairs 3. Glabrous Width/length of longest leaf. Ratio calyx length/length fruiting pedicle TI. 8. Ratio of flowering spike diameter, 3 mm from bottom of spike/ 3 mm from top. 9). 1 0. Capsule apex 0. Obtuse 1. Intermediate 2. Acute 11. Capsule length (mm) (lowest flower) 12. Seed number (average from five capsules) 13. Average length of five seeds (mm) TABLE 2. REGRESSION R* BETWEEN FREQUENCY OF JNTERMEDIA , DENSITY, INDEX, COEFFICIENT VARIATION OF MAJOR AND COEFFICIENT VARIATIONS (CV) Frequency intermedia Density both spp. cv major cv intermedia Frequency intermedia 1 Density both spp. ns 1 Cv major Ow Ole ns 1 cv intermedia 0-346* OSS 0-341 1 Unstained grains were considered unviable. The proportion of ovules setting apparently viable seed was examined by dissecting the contents of three capsules in a drop of water on a slide under a dissecting microscope. Infertile seeds were dark in colour, shrunken, and lacked mucilage. VARIATION IN SEED NUMBER AS A MEANS OF QUANTIFYING INTROGRESSION Thirteen 5 x 5 m areas were selected by eye from near the Metrocentre, and a single fruiting spike from every fruiting plant within each area was collected. The number of fruiting spikes per 25 m°* patch were counted, and the average seed number per capsule calculated for five capsules of each spike. Spikes were described as intermedia or major on the basis of average seed number per capsule. The discriminants used (major <10:4, intermedia 210-5) were based on previous findings in the literature, and a slight discontinuity observed in_ the distribution of the Metrocentre population data for this attribute. On the basis of this, the frequency of each taxon within the area was calculated. 376 R. EL-BAKATOUSHI, A. J. RICHARDS AND K. WOLFF For each area, two mothers were selected, one major and one intermedia (but in areas containing only major, two major mothers were selected) and 20 seedlings of each were grown to maturity (see seed germination) so that the coefficient of variation (cv) of seedling seed number could be calculated for each. This involved the raising of 2 x 13 x 20 = 520 seedlings. RESULTS MORPHOLOGY The first two components of the PCA represented 97:8 % and 1:9% of the total variation respectively. Of the 13 characters used (Table 1), six showed a significant correlation with both major principal components, and 10 showed a significant correlation with at least one. Three characters, all relating to fruits, did not significantly influence principal components, namely character 9 (relative length of fruting pedicel), 10 (capsule apex shape) and 11 (capsule length), although each of these showed some internal coherence with other characters used. The characters which mostly _ strongly influenced the two most important components were characters 2 (angle of blade to petiole) and 12 (seed number per capsule), two characters which traditionally have been important in the separation of the two taxa. Figure | represents the first two axes of PCA. The limits of variation in the standard populations are shown. The Metrocentre population encompassed the variation of the four ‘pure’ populations used as standards, and no less than 40% of individuals (18/45) fell outside the morphological limits of these standards, and can be considered to be morphologically intermediate between the taxa. Only three Metrocentre individuals corres- ponded clearly to intermedia, so that 24 individuals (53%) corresponded to major. No clear or distinct morphological clusters could be detected amongst or between Metrocentre populations, although the pure populations show distinct clustering. Most individuals of the intermediate phenotype falling outside the limits of the standard populations tended to resemble major more than intermedia, perhaps suggesting that most backcrossing occurs to major. HYBRID FERTILITY No significant difference was found in either seed set or pollen fertility between either parent and the offspring of crosses, or between the offspring of crosses and of selfs. Both the percentage of stainable pollen, and _ the proportion of ovules set as seed were invariably in excess of 80%. However, there was some evidence for significant matrilineal heritability in pollen stainability for selfed offspring from the Metrocentre population (1° = 11-6***), and for seed set after crosses in both Metrocentre and ‘pure’ populations (1° = 15-7* and r- = 46-5*** respectively). THE COEFFICIENT OF VARIATION OF MAJOR AND INTERMEDIA SEED NUMBER IN QUADRATS The regression between coefficient of variation (cv) of major seed number and frequency of intermedia per 25 m° quadrat showed that no ~ less than 70:1 % of the variability in seed number in major offspring could be attributed to the frequency of intermedia in that quadrat (Fig. 2 & Table 2). This suggested that considerable ongoing hybridisation occurred between major and intermedia. There was no significant effect between the density of fruiting plants per quadrat and offspring seed number cv (Table 2). However, the correlation between cv of intermedia offspring seed number and density of plants/quadrat was just significant (Table 2). There was also a positive relationship. between the frequency of intermedia and cv of seed number in intermedia offspring (t= 0:346*) (Table 2). The correlation between coefficient of variation of seed number of intermedia and cv of seed number of major was also slightly significant (r’ = 0:341*) (Table 2). DISCUSSION Using evidence from a variety of molecular markers, Morgan-Richards & Wolff (1999) suggested that major and intermedia are sufficiently distinct to warrant specific rank, despite evidence of widespread hybridisation outside the British Isles (Hooglander ef al. 1993, Wolff & Morgan-Richards 1999). However, the taxa are not so distinct in the British Isles (Kay 2002) and we doubt if morphologically ‘pure’ populations of inter- media often occur here. The scattered and largely ruderal distribution of intermedia in the INTROGRESSION BETWEEN PLANTAGO MAJOR SUBSPECIES 7) 0.35 - N 80%), seed set and fertility was high amongst manually made crosses, and the fertility of offspring of these crosses did not vary from that of their parents. Nevertheless, morphological differ- ences between the taxa in British populations persisted after cultivation, suggesting that many characteristics which separate the two taxa major and intermedia are genetically controlled (El-Bakatoushi 2004). In the Metrocentre population, intermediate, putatively hybrid individuals tended _ to resemble major from standard populations more than intermedia and this may be an indication that the hybrids have backcrossed to major individuals, which dominate in the community. (Another less likely hypothesis, i.e. that major characters are overwhelmingly dominant to intermedia characteristics was disproved after crosses between the taxa were reported by El-Bakatoushi (2004)). The highly significant relationship between variability in seed number in major offspring - and frequency of intermedia in the same quadrat suggested that this introgression is ongoing, reflecting a high rate of gene flow between two taxa and helped to confirm that some major individuals were introgressed towards intermedia. However, we also found a 378 R. EL-BAKATOUSHI, A. J. RICHARDS AND K. WOLFF 0.4 1.35 4 wo ©) oO cv of seed number offspring 0 0.1 0.2 0.3 0.4 0.5 0.6 relative frequency of intermedia in quadrat FIGURE 2 Scattergram between relative frequency of intermedia in quadrat and coefficient of variation of seed number of major (1° = 0-701), and intermedia offspring, showing lines of best fit. @ cv of seed number in major offspring, 0 cv of seed number in intermedia offspring positive relationship between coefficient of variation of seed number in intermedia and frequency of intermedia per quadrat. This could have been an artefact caused by a greater variability in seed number in intermedia than in major. The variability in intermedia seed number in general may have resulted from a greater tendency for intermedia individuals to receive outcrossed pollen from the numerically dominant plants of major. This suggestion was supported by the slightly positive relationship which occurred between density of plants per quadrat and the coefficient of variation for intermedia seed number. Minority effects occur when two potentially interfertile taxa or demes co-occur at very different frequencies, so that pollen arriving on stigmas of the minority taxon will originate overwhelmingly from the dominant taxon (Fowler & Levin 1984): Individuals of intermedia apparently suffered from minority effects at the Metrocentre which were maximised at high density, causing any outcrossed pollen to tend to originate from major. Consequently, one would expect immi- grant intermedia to lose their identity over successive generations, when it occurs, as here, in a minority Major and intermedia cannot be seen as separate species in the localities from which the mixed UK populations were collected, as it seems that genes of the infrequent and possibly adventive intermedia are readily diluted by the native taxon major. A high rate of gene flow occurred between these taxa, and both taxa could readily occupy the same open habitat. On this evidence the rank of subspecies is ideal for both taxa, even if the taxa sometimes behave more as one would expect of species in some non-British parts of their range. INTROGRESSION BETWEEN PLANTAGO MAJOR SUBSPECIES 37/9 ACKNOWLEDGMENTS We gratefully thank the University of Newcastle Fund for support to Ranya El-Bakatoushi. REFERENCES ABBOTT, R. J., ASHTON, P. A., & FORBES, D. G. (1992). Introgressive origin of the radiate groundsels, Senecio vulgaris L. var hibernicus Syme; Aat-3 evidence. Heredity 68: 425-435. ANDERSON, E. (1949). Introgressive Hybridization. Wiley, New Y ork. ANDERSON, E. (1953). Introgressive Hybridization. Biological Reviews 28: 280-307. ANDERSON, E. & HUBRICHT, L. (1938). Hybridisation in Tradescantia. Ill. The evidence for introgressive hybridization. American Journal of Botany 25: 396—402. CLAPHAM, A. R., TUTIN, T. G. & Moore, D. M. (1987). Flora of the British Isles, 3rd edition. Cambridge University Press, Cambridge. EL-BAKATOUSHI, R. (2004). Population genetics and evolution of the Plantago major group. Ph. D. thesis, University of Newcastle, UK. ESHED, Y. M., ABU-ABIED, Y. & ZAMIR, D. (1992). Lycopersicon esculentum lines containing small overlapping introgressions from L. pennellii. Theoretical and Applied Genetics 83: 1027-1034. FARRIS, M. A. & MITTON, J. B. (1984). Population density, outcrossing rate, and heterozygote superiority in Ponderosa pine. Evolution 38: 1151-1154. FOWLER, N. L. & LEVIN, D. A. (1984). Ecological constraints on the establishment of a novel polyploid in competition with its diploid progenitor. American Naturalist 124: 705-711. HARRISON, R. G. (1993). Hybrids and hybrid zones: a historical perspective, in R. G. HARRISON ed. Hybrid Zones and the Evolutionary Process, pp. 3-12. Oxford University Press, Oxford. HEGI, G. (1914). I/lustrierte Flora von Mitteleuropa. Vol. 6 (1) ed. 1. Munich. HOOGLANDER, N., LUMARET, R. & Bos, M. (1993). Inter- and intraspecific variation of chloroplast DNA of European Plantago spp. Heredity 70: 322-334. KAY, G. M. (2002). Plantago major subsp. intermedia, in C. D. PRESTON, D. A. PEARMAN & T. D. DINES eds. New Atlas of the British Flora, p. 535. Oxford University Press, Oxford. MORGAN-RICHARDS, M. & WOLFF, K. (1999). Genetic structure and differentiation of Plantago major reveals a pair of sympatric sister species. Molecular Ecology 8: 1027-1036. NEALE, D. B. & ADAMS, W. T. (1985). The mating system in natural and shelterwood stands of Douglas-Fir. Theoretical and Applied Genetics 71: 201-207. VAN DIk, H. (1984). Genetic variability in Plantago species in relation to their ecology. 2. Quantitative characters and allozyme loci in P. major. Theoretical and Applied Genetics 68: 43-52 VAN DIK, H. (1989). Genetic variability in Plantago species in relation to their ecology. 4. Ecotypic differentiation in P. major. Theoretical and Applied Genetics 77: 749-759. VAN DUK, H., WOLFF, K. & DE VRIES, A. (1988). Genetic variability in Plantago species in relation to their ecology. 3. Genetic structure of populations of P. major, P. lanceolata, and P. coronopus. Theoretical and Applied Genetics 75: 518-528. WOLFF, K. (1991). Analysis of allozyme variability in three Plantago species and a comparison to morphological variability. Theoretical and Applied Genetics 81: 119-126. WOLFF, K., FRISO, B. & VAN DAMME, J. M. M. (1988). Outcrossing rates and male sterility in natural populations of Plantago coronopus. Theoretical and Applied Genetics 76: 191-196. WOLFF, K. & MORGAN-RICHARDS, M. (1999). The use of RAPD data in the analysis of population genetic structure: case studies of Alkanna (Boraginaceae) and Plantago (Plantaginaceae), in P. M. HOLLINGSWORTH, R. M. BATEMAN & R. J. GORNALL eds. Molecular systematics and plant evolution,. pp. 51-73. Taylor & Francis, London. (Accepted December 2005) Watsonia 26: 381—383 (2007) Names and diagnoses for three willow (Salix L.) hybrids discovered recently in Scotland D. J. TENNANT Low Missise Farm, Laverton, Ripon, North Yorkshire HG4 3SY ABSTRACT Binomials and diagnoses are provided for three distinctive willow (Salix L.) hybrids found in recent years in the Scottish Highlands: Salix myrsinifolia x S. phylicifolia x S. repens = S. xlochsiensis; Salix arbuscula x S. repens = S. xlyonensis; and Salix caprea x S. myrsinifolia x S. phylicifolia = S. xmeikleana. INTRODUCTION Binomials and diagnoses are provided for three distinctive willow (Salix L.) hybrids which were discovered in recent years during field surveys of British montane willows, as summarised by Tennant (2001, 2004); full descriptions were provided by Tennant (2004). All three were discovered in montane habitats in the Scottish Highlands, but one of them (S. xmeikleana) has also been found in Mid-West Yorkshire (v.c. 64) (Tennant 2001, Wilcox 2001). They were all found growing near their parents and were all local and very scarce. S. xtetrapla Walker (S. myrsinifolia x S. phylicifolia) was probably one parent of S. xlochsiensis and of S. xmeikleana). Only two plants of S. xlochsiensis were found in Glenshee; one plant of S. xlyonensis was found on Meall Ghaordaidh, and two more near Creag Roro in 2002; and at least four plants of S. xmeikleana were found near Creag Roro. THREE NEW HYBRIDS Salix xlochsiensis D. J. Tennant, hybrida nova HOLOTYPUS: On stream-bank north-west of the Spittal of Glenshee, East Perthshire, v.c 89, GR NOO7, 5.8.1997, D. J. Tennant no. GSB 2H/97 (BM). Hybrida inter Salicem myrsinifoliam Salisb. et S. phylicifoliam L. et S. repentem L. A Salice myrsinifolia et S. phylicifolia et S. xtetrapla Walker (S. myrsinifolia x S. phylicifolia) habitu fruticis prostrati vel decumbentis nanissimi, ramos longos graciles gemmasque minores ferentis; foliis multo minoribus, marginibus revolutis undulatisque, primo dense sericeis, postea rarius sericeis, denique abaxialiter subglabris; petiolis saepe rubellis; amentis numerosioribus, minoribus, flores prorsum spectantes ferentibus, in pedunculis brevibus lateralibus _ portatis; amentorum squamis angustioribus, involventibus, omnino- viridibus_ vel nonnunquam dilute brunneis; ovariis capsulisque brevioribus distinguitur. Praeterea a §. myrsinifolia stipulis paucissimis vel absentibus et ovariis pubescentibus; a S. phylicifolia foliis serratis, dentes rubellos saepe ferentibus, abaxialiter venationem prominentem praebentibus, abaxialiter fere persistente sericeis muinusque glaucis, et pedicellis longioribus et stylis brevioribus distinguitur. A Salice repenti foliis magis undulatis, distinctissime serratis, dentes saepe curvatos acutos rubellos’ ferentibus, abaxialiter venationem prominentem praebentibus; amentorum squamis_ hirsutioribus, saepe anguste obovatis; stigmatibus longioribus distinguitur. A Salice xschraderiana Willd. (S. phylicifolia x S. repenti) foliis angustius obo- vatis (minus anguste ellipticis), obtusioribus, magis serratis, magis undulatis, dilute viridibus vel minus glaucis, abaxialiter venationem magis prominentem praebentibus; stipulis paucis nonnunquam praesentibus distinguitur. Hybrid between Salix myrsinifolia Salisb., S. phylicifolia L. and S. repens L. Distinguished from S. myrsinifolia, S. phylicifolia and S. xtetrapla Walker (S. myrsinifolia x S. phylicifolia) by being a prostrate or decumbent, very dwarf shrub with long, slender branches and smaller buds; by its leaves much smaller with revolute, undulate margins, initially densely sericeous, later more thinly sericeous, eventually subglabrous abaxially; by its petioles often reddish; by its catkins more numerous, smaller, with forward-directed 382 D. J. TENNANT flowers, and on short, lateral peduncles; by its catkin-scales narrower, involvent and wholly green or sometimes pale brown; and by its ovaries and capsules shorter. It is further distinguished from S. myrsinifolia by its stipules very few or absent and by its ovaries pubescent and from S. phylicifolia by its leaves serrate, often with reddish teeth, with prominent raised venation abaxially, almost persistently sericeous and less glaucous abaxially, by its pedicels longer, and by its styles shorter. Distinguished from S. repens by its more undulate leaves very distinctly serrate, with often curved, acute, reddish teeth and with prominent raised venation abaxially; by its catkin-scales more hairy and often narrowly obovate; and by its stigmas longer. Distinguished from S$ xschraderiana Willd. (S. phylicifolia x S. repens) by its leaves more narrowly obovate (less narrowly elliptic), more obtuse, more serrate, more undulate, and pale green or less glaucous with more prominent venation abaxially; and a few stipules sometimes present. Salix xlyonensis D. J. Tennant, hybrida nova HOLOTYPUs: On rock-ledge, Meall Ghaordaidh in Glen Lyon, Mid Perthshire, v.c. 88, GR NNS54, 1.9.1994, D. J. Tennant no. MG 2C/94 (BM). Hybrida inter Salicem arbusculam L. et S. repentem L. A Salice arbuscula’ foliis minoribus, hebetioribus, abaxialiter minus — glaucis, margines magis revolutos praebentibus, dentes pauciores minores_ ferentibus, adaxialiter persistentius appresso-pubescentibus, abaxialiter persistente vel dense sericeis; amentis squamas longiores ferentibus distinguitur. A Salice repenti foliis distincte serratis vel glanduloso-serratis, in sicco non nigrescentibus; amentis magis compactis, breviter pedunculatis, cum foliis primum visis, flores patentes (non prorsum_ spectantes) Squamasque latiores, ad apicem _ brunneas, densius pubescentes, ferentibus; ovariis densius pubescentibus, pedicello breviore, capsulam minorem parientibus, distinguitur. A Salice xpseudoglauca Andersson (S. arbuscula x §. lapponum L.) foliis minoribus, saturatius viridibus, nitidioribus, distinctius serratis, margines magis_ revolutos praebentibus, adaxialiter glabrioribus, abaxialiter pilos non nisi breves sericeos (non tomentosos) ferentibus; amentis minoribus, Ovaria capsulasque minores ferentibus, distinguitur. Hybrid between Salix arbuscula L. and S. repens L. Distinguished from S$. arbuscula by its leaves smaller, duller, iess glaucous abaxially, with more revolute margins, with fewer smaller teeth, more persistently appressed-pubescent adaxially, persistently sericeous to densely so abaxially; and by its catkins with longer catkin- scales. Distinguished from S. repens by its leaves distinctly serrate or glandular-serrate, not blackening when dried; by its catkins more compact, shortly pedunculate, appearing with | the leaves, with patent (not forward-directed) flowers, with wider, brown-tipped, more densely pubescent catkin-scales; and by its Ovaries more densely pubescent, with a shorter pedicel and producing a smaller capsule. Distinguished from S$. xpseudoglauca Andersson (S. arbuscula x S. lapponum) by its leaves smaller, deeper green, glossier, more distinctly serrate, with more revolute margins, more glabrous adaxially, with only short sericeous (not tomentose) hairs abaxially; and by its catkins smaller, with smaller ovaries and capsules. Salix xmeikleana D. J. Tennant, hybrida nova HOLOTYPUS: On _ stream-bank, Allt a’ Chobhair, Creag Roro, Glen Lyon, Mid Perthshire, v.c. 88, 7.7.2000, D. J. Tennant no. CR 9/00 (E). Hybrida inter Salicem capream L. et S. myrsinifoliam Salisb. et S. phylicifoliam L. A Salice caprea foliis adaxialiter vividius nitidiusque saturatiusque viridibus, abaxialiter plerumque glabris; ovariis brevioribus, pro ratione latioribus, stylum distinctum et stigmata vix longiora ferentibus, distinguitur. | A Salice myrsinifolia fruticis plerumque excelsioris habitu; foliis latioribus, magis coriaceis, adaxialiter vividius nitidiusque plerumque _ saturatius viridibus, minus pubescentibus, in sicco non nigrescentibus; ovariis uniformiter pubescentibus, stigmata vix longiora ferentibus, distinguitur. A Salice phylicifolia ramunculis —lentius glabrescentibus; foliis magis serratis (quorum NAMES AND DIAGNOSES FOR THREE SALIX HYBRIDS 383 maxima ad apicem basinque latius rotundata sunt), apicem versus et adaxialiter in costa persistente sed tenuiter pubescentibus, abaxialiter venationem magis prominentem praebentibus; stipulis plerumque bene effectis; pedicellis longioribus ubi capsulae effectae sunt; stylis paulo brevioribus distinguitur. A Salice xlatifolia Forbes (S. caprea x S. myrsinifolia) foliis maximis plerumque latioribus, ad apicem obtusioribus, magis cori- aceis, adaxialiter vividius nitidiusque satura- tiusque vviridibus, abaxialiter plerumque glabris, in sicco non _ nigrescentibus, distinguitur. Ab hybrida Salice caprea x S. phylicifolia foliis apicem versus et adaxialiter in costa persistentius sed tenuiter pubescentibus, abax- ialiter plerumque_ glabris; stipulis saepe majoribus distinguitur. A Salice xtetrapla Walker (S. myrsinifolia x S. phylicifolia) praecipue gemmis majoribus, luteolis; foliis maximis pro ratione latioribus, ad apicem basinque latius rotundatis, in sicco non partim nigrescentibus; stipulis saepe magis effectis distinguitur. Hybrid between Salix caprea L., S. myrsinifolia Salisb. and S. phylicifolia L. Distinguished from S. caprea by its leaves brighter, glossier, deeper green adaxially and usually glabrous abaxially; and by its ovaries shorter and relatively broader, with a distinct style and marginally longer stigmas. Distinguished from S. myrsinifolia by being a usually taller bush; by its leaves wider, more coriaceous and brighter, glossier, usually deeper green adaxially, less pubescent, and not blackening when dried; and by its ovaries uniformly pubescent with marginally longer stigmas. Distinguished from S. phylicifolia by its twigs more slowly becoming glabrous; by its leaves more serrate and persistently but thinly pubescent near the apex and on the midrib adaxially, with more prominent venation abax- lally, the largest leaves more broadly rounded at the apex and base; by its stipules usually well developed; by its pedicels longer when capsules have developed; and by its styles slightly shorter. Distinguished from S. x latifolia Forbes (S. caprea X S. myrsinifolia) by its largest leaves usually broader, more obtuse at the apex, more coriaceous, brighter, glossier, deeper green adaxially, usually glabrous abaxially, and not blackening when dried. Distinguished from S&S. caprea x _ S. phylicifolia by its leaves more persistently but thinly pubescent near the apex and on the midrib adaxially and _ usually glabrous abaxially; and by its often larger stipules. Distinguished from S. xtetrapla Walker (S. myrsinifolia x S. phylicifolia) mainly by its buds larger and yellowish; by its largest leaves relatively broader, more broadly rounded at the apex and base, and not blackening in parts when dried; and by its stipules often more developed. This hybrid is named after Robert Desmond Meikle in recognition of his long service and considerable contribution to the study of British willows. ACKNOWLEDGMENTS I would like to thank Desmond Meikle for his confirmation or initial determination of the hybrids, Clive Stace and Philip Oswald for their guidance and helpful suggestions for the preparation of this note, and Philip Oswald for the Latin translation. REFERENCES TENNANT, D. J. (2001). Salix caprea L. x S. myrsinifolia Salisb. x S. phylicifolia L. in Perthshire. Watsonia 23: 547-549. TENNANT, D. J. (2004). A re-assessment of montane willow (Salix L., Salicaceae) hybrids in Scotland. Watsonia 25: 65-82. WILCOX, M. (2001). Two triple hybrid willows in the Yorkshire Dales. BSB News 86: 28-29. (Accepted August 2006) ill | i ce ‘ 4 | i ay ‘ ” H ’ | | i ) - i - x j Heep . ) f ? | ‘ | > p } E ns | ier i n Watsonia 26: 385—389 (2007) Notes THE GUNNA (V.C. 103) RECORD FOR CAREX APPROPINQUATA SCHUM. (CYPERACEAE) Many of the late 1930s and 1940s records of J. W. Heslop Harrison from the Isle of Rhum (v.c. 103) and elsewhere in the Hebrides are now widely considered to result from the deliberate introduction of plants in an attempt to provide evidence to support his theory for survival of elements of the flora from pre- glacial times (Sabagh 2001, Preston 2004, Pearman & Walker 2004). Several such records are for Carex species, some of them purporting to be the first for the British Isles. Perhaps the most surprising were Carex bicolor and C. glacialis both known from Europe and elsewhere but not from the British Isles. Understandably, these have been treated with considerable scepticism, both at the time and since. There is, however, a most interesting Heslop Harrison record for Carex appropinquata from the islet of Gunna (v.c. 103) in the Inner Hebrides which lies between Coll and Tiree. This record was first published under the name C. paradoxa Willd. in The Flora of the Isles of Coll, Tiree and Gunna (Heslop Harrison et al.1941) with the statement: “Rare in Gunna, and only recorded once previously from a Scottish locality. This was in Peebles”. If correct, this would represent a considerable extension of range since the stronghold for C. appropinquata in the British Isles is in East Anglia; elsewhere it is only known locally in Yorkshire, the Scottish Borders and Ireland. Whilst examining Carices in the Edinburgh herbarium (E), it was surprising to find a previously overlooked voucher specimen which supports this record. The specimen is of a rather immature plant but which, by its general facies and its narrow leaves (to a maximum width of 2mm) and black fibrous basal leaf sheaths, is undoubtedly C. appropinquata. Indeed there is an added determination slip to that effect signed by Ernest Nelmes, the Kew specialist, to whom Heslop Harrison often sent material for confirmation. The specimen has been in the Edinburgh herbarium for over twenty years and was formerly in the collection of Robert Mackechnie, a colleague of Heslop Harrison who was invited by the latter to botanise with him on Rhum in _ 1949 (Mackechnie in litt. to A. C. Jermy). This specimen, along with others, was bequeathed to the Edinburgh herbarium shortly after Mackechnie died in 1978. The label states “Carex appropinquata. Island of Gunna, between Coll & Tiree. June 1940. Prof. J. W. Heslop Harrison”. However, the label is not in Heslop Harrison’s hand but was directly transcribed by the curator from the wrapping covering the specimen (which also contained Nelmes’ original signed deter- mination slip) when received for incorporation (D. R. McKean pers. comm. 2006). The curator feels confident that this was done accurately. There are certain facts which support this record and others which cast doubt upon it. Perhaps the most favourable is that it is a published record supported by a _ voucher specimen, albeit on a re-labelled sheet. Also, a Hebridean record for C. appropinquata is unlikely to have been of value in support of Heslop Harrison’s theory relating to pre-glacial survival and, therefore, to have been deliberately planted by him (or even by an associate). Although there had been an old unconfirmed record from Innerleithen, Peeblesshire (v.c. 78), by Lyell in the 1850s (David 1990), subsequent to the 1940 Gunna record, C. appropinquata was not re-found in that general (Borders) area of Scotland until found in Roxburghshire (v.c. 80) in 1967 (Corner 1969). Heslop Harrison was obviously aware of the old record (Heslop Harrison et al. 1941) but it seems very unlikely that his specimen could have originated from there. In the British Isles, C. appropinquata has a rather disjunct distribution pattern, so that its occurrence in the Inner Hebrides is quite possible. Also, C. paniculata and C. diandra, two closely related species, the latter especially being a close ecological associate of C. appropinquata as at Malham Tarn (v.c. 64), both occur on neighbouring Tiree (Pearman & Preston 2000). It is also known that Heslop Harrison’s party explored Gunna in the summer of 1940 and it was in June of that year that the specimen was collected. Due to the suspicion surrounding many of Heslop Harrison’s 386 NOTES Watsonia 26 (2007) records, it 1s tempting to dismiss anything unusual as being of dubious worth but this is perhaps unfair since there are some of his Hebridean rarities which have been sub- sequently confirmed, e.g. Spiranthes rom- anzoffiana. There is also his collection of the Asiatic Carex brunnea Thunb. from the grounds of Kinloch Castle (Rhum), again confirmed by Nelmes, which he readily conceded (Heslop Harrison 1945 and in sched.) as having been almost certainly introduced accidentally with planted bamboo. The voucher specimen for this is also in the Edinburgh herbarium (E) and again originated via Mackechnie in the same way as that of the Gunna specimen of C. appropinquata. An annotation on the C. brunnea sheet in Heslop Harrison’s hand states that it was collected on August 12, 1944. On the other hand certain factors cast doubt on the Gunna record. Firstly, it is surprising that it is unmentioned by Heslop Harrison (1948) in his later review of his own records of noteworthy sedges from the Inner and Outer Hebrides, although he does ‘include its close associate C. paniculata for neighbouring Tiree. In addition, Pearman & Preston (2000) discount the record of C. appropinquata on ecological grounds stating that “There is no suitable habitat on Gunna for this species”’. It is also possible that the collection may not have been made by Heslop Harrison in person since he states (Heslop Harrison et al 1941) that in the years 1939 and 1940, the co-authors of that paper (although perhaps not necessarily always present himself) spent three long periods on Coll and Tiree. On the second of these periods (whether 1939 or 1940 is not clear) whilst based on Tiree they also explored Gunna. However in 1939 a party of his students also camped on Coll and “broke new ground on the Isle of Gunna’. Perhaps it was the students who were thought, or claimed, to have coll- ected the C. appropinquata or, dare one suggest, attempted to deceive by introducing it there? Unless C. appropinquata is re-discovered on Gunna, the record is likely to remain a mystery. Despite suspicion over Heslop Harrison’s more unusual records, it is difficult to see how a falsification would have any relevance in ~ furthering his theory of Hebridean plant survival from pre-glacial times. The alternative conclusion is that there has been a mis- labelling of the specimen or other mix-up when collecting. ACKNOWLEDGMENTS I am grateful to A. C. Jermy for allowing me to examine Mackechnie’s letter and to D. R. McKean for information on the provenance of the Gunna specimen. REFERENCES CORNER, R. W. M. (1969). Carex appropinquata Schumach. — In Scotland. Plant Notes. Proceedings of the Botanical Society of the British Isles, 7: 562. DAVID, R. W. (1990). The distribution of Carex appropinquata Schumacher (C. paradoxa Willd.) in Great Britain and Ireland. Watsonia 18:201-—204. HESLOP HARRISON, J. W. (1941). The Flora of the Isles of Coll, Tiree and Gunna (v.c. 110B). Proceedings of the University of Durham Philosophical Society, 10(4): 301-304. HESLOP HARRISON, J. W. (1945). Noteworthy Sedges from the Inner and Outer Hebrides, with an Account of Two Species New to the British Isles. Transactions and Proceedings of the Botanical Society of Edinburgh, 270-277. PEARMAN, D. A. & PRESTON, C. D. (2000). A Flora of Tiree, Gunna & Coll, p. 68. Dorchester. PEARMAN, D. A. & WALKER, K. J. (2004). An examination of J. W. Heslop Harrison’s unconfirmed plant records from Rum. Watsonia 25: 45-63. PRESTON, C. D. (ed.) (2004). John Raven’s Report on his visit to the Hebrides, 1948. Watsonia, 25: 17-44. SABBAGH, K. (2001). A Rum Affair. A True Story of Botanical Fraud. 1-276. Da Capo Press. M. J. Y. FOLEY Faraday Building, Dept. of Biological Sciences, University of Lancaster, Lancaster LAI 4YA NOTES Watsonia 26 (2007) 387 VALERIANELLA ERIOCARPA DESV. AS A PRESUMED NATIVE IN BRITAIN — AN UPDATE In our paper in Watsonia (Pearman & Edwards 2002), we suggested that V. eriocarpa was a native member of an annual early flowering cliff-edge community on the limestones of Dorset and the hard chalk of the Isle of Wight. We had also looked, without success, on the South Devon limestone at Berry Head, and felt ambivalent about the status of the relatively persistent Cornish records, which are on sand. Since then there have been several interest- ing developments, with records from completely new areas. VC3, SOUTH DEVON. Ilsham Marine drive, Torquay. SX941632. A. J. Byfield. 2002. A few plants in short therophyte turf on steep S facing road verge bank. Headland north of Whitsand Beach, Maidencombe. SX927676. D. Buckingham. <1995. A few plants in a relict fragment of unimproved cliff-top grassland. Not searched for again. VC49, CAERNARVONSHIRE. Great Orme, Llandudno. SH773824, (c. 20 plants in 2003), SH772827, (c. 30 plants in 2003), SH776827, (c. 45 plants in 2003). W. McCarthy. 1992 and every year since. On very thin soils over outcropping limestone, south-facing, with a range of species including Aphanes arvensis, Arenaria serpyllifolia, Carex caryophyllea, Cerastium pumilum, C. semidecandrum, Erophila glabrescens, Festuca ovina, F. rubra, Galium verum, Sherardia arvensis, Thymus polytrichus, Veronica arvensis. These new sites then are limestone head- lands, with broadly similar communities to the Dorset sites. In addition, one of us (DAP) searched the Gower limestones for two days in May 2005 without any success, finding only V. locusta, and that only in very small quantity. This means that of all the coastal limestone areas in southern England and Wales, only those at Weston-super-Mare and Brean Down (VC6) remain to be searched. Elsewhere Ted Pratt and David Leadbetter have added some new sites in Dorset, slightly further inland on the Purbeck limestone and at Corfe Castle on the chalk. These are from permanent grassland, albeit over rock and with Open areas. Conversely new areas of arable there have produced Euphorbia platyphyllos and V. dentata but no V. eriocarpa. DAP has visited the two Cornish sites mentioned in the Watsonia article (Pearman & Edwards 2002). At the Constantine site (SW8675) the sandy wall had become over-grown (though it has just been cleared), and no plants have been seen there for 15 years, but at the second site, at Phillack Towans, near Hayle (SW5538) there are plenty of patches. But here it looks so ruderal, growing on walls, waste lots and around buildings, that it is difficult to see it as a native. A further and more natural site has been discovered by Ian Bennallick below the coast path at Harbour cove, Padstow (SW9177) where it grows on rocks just by the beach with V. rimosa nearby! The coastal footpath separates this site from arable fields above, with Vicia bithynica on the edge, so the site has more of the flavour of an arable weed refugium. We still feel uncertain of the status of the Cornish sites, though we tend even more to considering them as persistent aliens, but the records from the new limestone sites are potentially very interesting indeed. We are very grateful to those named above for the new records and details. REFERENCE PEARMAN, D. A. & EDWARDS, B. (2002). Valerianella eriocarpa Desv. in Dorset, and a reassessment of its status as a as a presumed introduction in Britain. Watsonia 24: 81-89. D. A. PEARMAN Algiers, Feock, Truro, Cornwall, TR3 6RA B. EDWARDS Bere Heath Farm, Bere Regis, Dorset, BH20 7NS 388 NOTES Watsonia 26 (2007) RUBUS MILESIANUS WHOLLY A GARDEN ESCAPE IN BRITAIN? Certain members of the Rubus fruticosus aggregate characterised by fruit of a kind holding a special appeal to growers have probably long been liable to be taken into cultivation from the wild and in some cases marketed commercially. Unfortunately, it has not been the practice in the past for examples of what nurseries had on offer to be preserved in herbaria: field botanists have collected cultivated taxa only when those have chanced to occur outside the confines of gardens sufficiently to be claimable as ‘wild’. A rare documented exception is R. bartonii Newton, a member of Series Vestiti widespread as an unquestioned native in Wales and western parts of England with a marked peak of abundance in Cardiganshire, v.c. 46 (in which it is the commonest bramble). Cultivated at the former Long Ashton Research Station, near Bristol, and some years ago placed on the market under the trade name ‘Ashton Cross’ (Edees & Newton 1988: 127 fn.), this has won favour comparatively widely by reason of its heavy cropping despite rather acidic fruit. Isolated occurrences of it well outside its presumptive natural range, as on Barnes Common in Surrey, v.c. 17 (Norman 1999), can be credibly attributed to garden outcasts or dispersal by birds from cultivated sources; elsewhere, though, some supposedly native populations of the species may be more suspect in status than is at first sight apparent. In the Isle of Man, for example, in the one place in which it occurs in quantity, in a shady ravine Opening on to the sea, a habitat which seemed good grounds for regarding it as a member of the native flora (Allen 1986), the population has subsequently been found to extend into an adjacent garden, the owner of which turns out to have long prized this particular bramble for the very qualities for which it has been promoted commercially. In the light of this finding, the status of this species in the island as a whole has come into question. A more extreme possibility is that some supposedly native species are garden escapes in Britain (and Ireland too, for that matter) in their entirety. One of those, it has belatedly become apparent, could well be the recently-described R. milesianus. Robust and distinctive enough to have gained a place in the herbaria of Rubus specialists since as early as 1867 (Watson 1958: 194), its recorded occurrences all lie along an arc of some 235 km in length extending across the far south-east corner of England, from Bournemouth to Canterbury. Most of these appear to have taken the form of solitary bushes only, a pattern especially characteristic of larger-fruited species such as this that are consequently more than ordinarily prone to dispersal by birds — though in two instances secondary spread by tip-rooting has clearly taken place on an extensive scale. In addition to their large size the mature fruits of R. milesianus are also notably delicious. It is thus a species particularly likely to have been taken into cultivation from the wild. The possibility that is may have occurred as a garden escape has nevertheless been raised hitherto only in the case of one anomalous- — looking urban record (Allen 2004: 170). Subsequent scrutiny of all the recorded occurrences, however, has revealed a suggest- ive association with areas notable for a concentration, at least by the mid-nineteenth century, of large detached houses standing in grounds spacious enough to have contained a sizeable kitchen-garden extending to a range of soft fruit. Even one of the two populations of considerable extent, that on Southampton Common, v.c.11, is suspiciously confined to the neighbourhood of the Common’s north margin, the part adjacent to that city’s leafy, late-developing suburb of Bassett. At first sight the other substantial population, by far the larger of the two in extending across 3 x 2 km of wooded countryside in north-east Hampshire, v.c.12, has no comparably suspect character. Although that area has its south end as the type locality — the extensive grounds of a mansion, Tylney House (now a hotel) — the readiest assumption is that the plant owes its presence in those to overspill from the adjoining woodland. Equally, however, the reverse could be the case. Given the impress- ively rapid rate of spread by tip-rooting of which Rubus species are well-known to be capable, it would not have been difficult for one as robust as this to have colonised that large tract of country within a relatively short span of years. The more or less continuous character of the population could be seen as providing support for that alternative interpretation. But if the status of R. milesianus in Britain is wholly that of a naturalised escape, there remains the question of from where it NOTES Watsonia 26 (2007) 389 originated. Did it arise in cultivation (as the widely-grown R. laciniatus Willd. is supposed to have done, in the absence of contrary evidence) or does it exist somewhere as a native? After a century and a half in which Britain has been’ explored increasingly intensively by Rubus specialists, it is hard to believe that so conspicuous a species still lies undiscovered in such situations here. Mainland Europe accordingly seems much likelier as the source. And to where more precisely that might have been, the recent surfacing in BM of an unmounted gathering from southern France of what has every appearance of being a weak example of this same taxon provides a clue. These specimens were collected in 1925 by a then leading Swedish batologist, C. E. Gustafsson, in or near Carmaux, a town in the north of dép. Tarn, in the south-western foothills of the Massif Central. They are accompanied by a label in the handwriting of the collector with a determination that he probably arrived at by himself from the keys and descriptions in Rubi Europae (Sudre 1908- 1913): R. micans subsp. heterochrous var. nitidipilus Sudre, a taxon described in that work from another locality in that same départemente. As no authentic material of that has been located, it is impossible to say whether that determination is correct (which could have nomenclatural implications, if it is ). The specimen does, however, provide a possible homeland for R. milesianus — assuming that the plant found by Gustafsson was not an escape there as well. In addition it reopens the possibility that Watson (1958) may have been correct after all in identifying the British plant with R. koehleri subsp. lapeyrousianus Sudre, a taxon described (and known only) from dép. Ariége, at the east end of the Pyrenees. Though Arie¢ge is two départementes distant from Tarn and situated alongside a different mountain range, the separating distance is not so great as to make such an identification geographically altogether unlikely. However, that question too cannot be settled for similar lack of an authentic specimen (Allen 2004). REFERENCES ALLEN, D. E. (2004). Five new species of Rubus L. (Rosaceae) mostly from Central South England. Watsonia 25: 157-174. EDEES, E. S. & NEWTON, A. (1988). Brambles of the British Isles. Ray Society, London. NORMAN, E. (1999). The flora of Barnes Common. London Naturalist 78: 65—80. SUDRE, H. (1902-1913). Rubi Europae. Librairie des Sciences naturelles, Paris. WATSON, W. C. R. (1958). Handbook of the Rubi of Great Britain and Ireland, ed. P. D. SELL & J. E. WOODHEAD. Cambridge University Press, Cambridge. D. E. ALLEN Lesney Cottage, Middle Road, Winchester, Hampshire, SO22 5EJ HI Wii) Mi Nel | Hi i Heel i] Whe (Nt i) j Ht i} | } WW i | H) I {Wa nit | wh i Watsonia 26: 391—407 (2007) 391 Plant Records Records for publication must be submitted to the appropriate Vice-county Recorder (see BSBI Year Book 2005), and not to the Editors. Following publication of the New Atlas of the British & Irish Flora and the Vice-county Census Catalogue, new criteria have been drawn up for the inclusion of records in Plant Records. (See BSBI News no. 95, January 2004 pp10 & 11). These are outlined below: e First records of all taxa (species, subspecies and hybrids) included in the VCCC, designated as native, archaeophyte, neophyte or casual. e First record since 1970 of the taxa above, except in the case of Rubus, Hieracium and Taraxacum. e Records demonstrating the rediscovery of all taxa published as extinct in the VCCC or subsequently. e Newly reported definite extinctions. e Deletions from the VCCC (e.g. through the discovery of errors, the redetermination of specimens etc.) NB — only those errors affecting VCCC entry. e New 10km square records for Rare and Scarce plants, defined as those species in the New Atlas mapped in the British Isles in 100 10km squares or fewer. (See BSBI News no. 95, January 2004 pp 36-43). Records for the subdivisions of vice-counties will not be treated separately; they must therefore be records for the vice-county as a whole. However, records will be accepted for the major islands in v.cc. 102—104, 110 and 113. In the following list, records are arranged in the order given in the List of Vascular Plants of the British Isles and its supplements by D. H. Kent (1992), from which the species’ numbers, taxonomy and nomenclature are taken. The Ordnance Survey national grid reference follows the habitat and locality. With the exception of collectors’ initials, herbarium abbreviations are those used in British and Irish Herbaria by D. H. Kent & D. E. Allen (1984). Records are field records if no other source is stated. For all records ‘det.’ or ‘conf.’ appear after the herbarium if the determination was based on material already in an institutional herbarium, otherwise before the herbarium. The following signs are used: * before the vice-county number: to indicate a new vice-county record. + before the species number: to indicate that the plant is an archaeophyte. + before the species number: to indicate that the plant is a neophyte. © before the species number: to indicate that the plant is a casual. The above 3 signs may also used before the vice-county number to-indicate the status of the plant in that vice-county. ® before the vice-county number: to indicate that this is an additional hectad for a Rare or Scarce plant. © at end of entry: established taxon not in Vice-County Census Catalogue. [ ] enclosing a previously published record: to indicate that that record should be deleted or changed. The following list contains the second set of records up to and including the year 2005. Records up to and including the year 2006 will be published in the next issue of Watsonia. 1/3.1. Lycopodium clavatum 110, Outer Hebrides: Lingara, South Harris, NGO0585, R. J. - Pankhurst, R. A. H. Smith & E. Pilling, 1997. Ist record since 1970. 4/1.2. Equisetum ramosissimum *35, Mons.: in rough grass, a brownfield site, NW of Spytty Pill, Newport, ST323874, R. P. Kilshaw, 2005, conf. T. G. Evans, A. C. Jermy, P. J. Acock & F. J. Rumsey. Ist Welsh record. Translocation is being considered. 4/1.3. Equisetum variegatum 106, E. Ross: forested dunes, Morrich More, NH802833, B. R. & C. B. Ballinger, 2005, herb. B.B. 1st record since 1970. 392 PLANT RECORDS 4/1.4x8.Equisetum xdycei (E. fluviatile x E. palustre) *93, N. Aberdeen: flush, Scad Hill, NJ428245, P. J. Acock, F. J. Rumsey & D. Welch, 2003, herb. D. Welch. Known in Scad Hill - Silverford flushes since c. 1980, but identity uncertain. 4/1.6. Equisetum pratense 110, Outer Hebrides: Rodel, Harris, NG041845, C. N. Page, 1972, E. 1st record since 1970. 4/1.6x7. Equisetum xmildeanum (E. pratense x E. sylvaticum) *93, N. Aberdeen: flush in gorge, White Hill of Bogs, NJ438253, P. J. Acock, F. J. Rumsey & D. Welch, 2005, herb. D. Welch. E. xmildeanum suspected in 2003, confirmed at same position in 2005. 6/1.1. Osmunda regalis 95, Moray: one plant in damp area under pine trees, Lossie Forest, NJ314658, I. P. Green, 2005. Ist record since 1955. +8/1.nip. Pteris nipponica Shich *29, Cambs.: a single naturalised plant near base of wall, Essex Building, Queens’ College, Silver Street, Cambridge, TL446581, P. H. Oswald, 2000. @ 11/1.1x2. Polypodium xmantoniae (P. vulgare x P. interjectum) *110, Outer Hebrides: Abhainn Gheurasdail, South Harris, NG107969, P. A. Smith, 2005, conf. G. Hutchinson, NMW. 14/1.1. Thelypteris palustris ®28, W. Norfolk: fen, Sculthorpe, TF902300, R. A. Yaxley, 2001. @®28, W. Norfolk: fen, Sculthorpe, TF901297, R. A. Yaxley, 2001. ®28, W. Norfolk: fen grassland, East Walton Common, TF734166, Norfolk Flora Group & R. W. Ellis, 2002. 14/3.1. Oreopteris limbosperma — 32, Northants.: one plant only growing near patch of ling by old railway tunnel, Charwelton, SP533569, J. Atkinson, 2004, conf. I. A. Denholm, herb. Gill Gent. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995). 15/2.5b. Asplenium trichomanes subsp. quadrivalens *91, Kincardines.: roadside dyke, Banchory, NO693959, D. Welch, 2005, herb. D. Welch. Site long known but VCCC recorded only the species, not the subspecies. 15/2.7. Asplenium ruta-muraria 106, E. Ross: urban site, Muir of Ord, NH5250, M. J. Crawley, 1998. Ist record since 1970. Still present in Muir of Ord 2001. 16/5.1. Cystopteris fragilis £32, Northants.: wall, Brockhall, SP632626, B. J. Laney, 2005, conf. P. Stanley, herb. Gill Gent. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995). 17/1.1. Polystichum setiferum 104, N. Ebudes (Skye): cliff base, S of Holm, NG519507, S. J. Bungard, 2004, conf. M. H. Rickard. Ist record for Skye. **110, Outer Hebrides: old woodland garden, Northbay, Barra, NF705036, R. J. Pankhurst, 2004, det. H. McHaffie, E. 17/1.2. Polystichum aculeatum 110, Outer Hebrides: ravine NE of Clett an Duin, Taransay, NB037003, E. Pilling, 1997, E. Ist record since 1970. 17/3.1. Dryopteris oreades *110, Outer Hebrides: ravine of Abhainn a’ Mhuill, NB188130, R. J. Pankhurst, 1996, det. S. Lindsay, E. Ist confirmed record. 17/3.2x3. Dryopteris xcomplexa (D. filix-mas x D. affinis) *8, S. Wilts.: woodland, Great Bedwyn, SU283626, Wiltshire Botanical Society, 2003. 17/3.2x3. Dryopteris xcomplexa nothosubsp. critica (D. filix-mas x D. affinis subsp. borreri) *46, Cards.: wooded ravine, Llyfnant Valley, 300m SW of Cwmrhaiadr, SN752960, T. D. Dines & A. O. Chater, 1997, det. K. Trewren, NMW. 17/3.3con. Dryopteris affinis morphotype convexa *44, Carms.: acid boundary bank, Glandy Farm, SN167287, BSBI meeting, 2005, det. A. O. Chater & S. J. Thomas. 1st record of morphotype. 17/3.3pal. Dryopteris affinis morphotype paleaceo-lobata *44, Carms.: hedgebank, Tireithin, Bancffosfelen, SN493127, BSBI meeting, 2003, det. A. O. Chater & S. J. Thomas. Ist record of morphotype. +20/7.mug. Pinus mugo 110, Outer Hebrides: Glen Coishletter, South Harris, NF998885, R. J. Pankhurst, 2002. Ist record since 1970. t20A/CRY.jap. Cryptomeria japonica *7, N. Wilts.: two seedlings progressing to saplings in woodland, Chisbury Wood, SU275655, J. E. Oliver, 2004. +21/1x2/1xnoo. xCupressocyparis leylandii (Cupressus macrocarpa xChamaecyparis nootkatensis) *110, Outer Hebrides: Scotasay, NG188972, R. J. & A. F. Pankhurst, 2005. ©28/3.3. Helleborus orientalis *7, N. Wilts.: established on dumped soil, Clatford, SU160688, J. E. Oliver, 2003. *28/3.arg. Helleborus argutifolius *©5, S. Somerset: self-sown at base of pavement, Witcombe, ST476214, P. R. Green, 2005. © PLANT RECORDS 393 +£28/12.4. Clematis montana *©29, Cambs.: one self sown plant in paving crack below Elizabeth Way bridge, Cambridge, TL461591, A. C. Leslie, 2005. 28/13.4. Ranunculus sardous 32, Northants.: field edge, SP663658, B. J. Laney, 2005, det. E. J. Clement. Ist record since 1970. 28/13.13x14. Ranunculus xlevenensis (R. flammula x R. reptans) *108, W. Sutherland: at 553m, Coir a’ Ghallaich, Ben Hope, NC486505, G. P. Rothero, 2004, conf. R. J. Gornall. 28/13.23. Ranunculus aquaticus 106, E. Ross: farm pond near Rockfield, NH930838, B. R. & C. B. Ballinger, 2005. Ist record since 1970, confirming Atlas record for NH98, M. J. Crawley, 995% ©29/1.7. Berberis julianae *7, N. Wilts.: roadside, Marlborough, SU192695, J. E. Oliver, 2004. Recent garden escape. +29/1.9. Berberis darwinii *7, N. Wilts.: five self-seeded young plants on stonework, Chippenham, ST920735, J..E. Oliver & M. Blake, 2003. *51, Flints.: on established coastal dunes away from houses, Talacre, SJ1285, J. Phillips, 2005. +30/1.3. Papaver somniferum *+110, Outer Hebrides: Eoropie, NB519649, R. J. Pankhurst, 2005. +30/1.6. Papaver hybridum 32, Northants.: arable field edge, Spanhoe, SP945987, A. Wilson, 1999, det. R. J. Wilson. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995) and Ist record since Druce 1930. +31/3.1. Pseudofumaria lutea 106, E. Ross: wall, Rosemarkie, NH736577, B. R. & C. B. Ballinger, 2005. Ist record since 1970. 31/4.1. Ceratocapnos claviculata *32, Northants.: very well established in conifer plantation, Harlestone Firs, SP718640, K. Taylor, 1996, conf. G. M. Gent, herb. Gill Gent. 31/5.1. Fumaria capreolata 50, Denbs.: footpath, Colwyn Bay, SH875787, J. A. Green, 2005. Ist record since 1960. +31/5.10. Fumaria vaillantii ®28, W. Norfolk: urban area, Dereham, TF999131, D. & B. Mathias, 2004. +£32/1.occxori. Platanus xhispanica (P. occidentalis x P. orientalis) *5(0), Denbs.: streamside, Alyn Waters County Park, Wrexham, SJ319549, BSBI group, 2005, NMW. +36/1.1b. Urtica dioica subsp. galeopsifolia *44, Carms.: rubbish tip, old quarry north of Beacon Bog, SN31N, C. Cheffings, M. Godfrey & A. R. Vickery, 2005. Ist Welsh record for subspecies 40/1.2b. Betula pubescens subsp. tortuosa *110, Outer Hebrides: Molinginish, NB2200, J. W. Heslop Harrison, 1956. Heslop Harrison et al in Proc. Unic. Durh. Phil. Soc. 12: 141-149 (1956). 40/1.3. Betula nana = *86, Stirlings.: Craigend Muir, Strathblane, NS581774, J. Mitchell & D. C. Shenton, 2002, E. t40/1.utixl. Betula utilis x B. pendula *29, Cambs.: one self sown tree, apparently of this parentage, on a grave at the southern end of Mill Road cemetery, Cambridge, TL461561, A. C. Leslie, 2001, CGE. First seen in 2001 when 6ft tall. Still there’ (2005) and now 15ft tall. © Apparently Ist record for the British Isles. t40/2.1x2. Alnus xhybrida (A. glutinosa x A. incana) *44° Carms.: self sown, with both parents in vicinity, amongst shrub plantings, Wildfowl & Wetlands Trust Centre, Penclacwydd, Llanelli, SS533986, Llanelli Naturalists, 1999, det. I. K. Morgan. *©57, Derbyshire: amenity tree planting, near Grin Low car park, SK049719, A. Willmot & J. Bland, 2004. Probably planted, and with both parents which were planted. +40/2.3. Alnus cordata *7, N. Wilts.: seedlings and 3 saplings on canalside, Swindon, SU135836, J. E. Oliver, 2004. £40/2.rub. AInus rubra *42, Brecs.: Glasfynydd Plantation, SN82, R. G. Woods, 2004. 743/1.12. Chenopodium murale ©46, Cards.: several plants on waste ground, Ynys-las _ boatbuilding yard, SN616932, A. O. Chater, 2005, NMW. Ist record since 1932. ©43/1.gig. Chenopodium giganteum *29, Cambs.: one plant on a low ridge of waste soil, Jeavons Lane, Cambridge, TL320588, N. P. Millar, 2005, CGE. 43/3.3x4. Atriplex xtaschereaui (A. glabriuscula x A. longipes) *95, Moray: several plants in saltmarsh on the edge of Lossie Forest, Lossiemouth, NJ253689, I. P. Green, 2005. 394 PLANT RECORDS 43/3.6. Atriplex litoralis | *©32, Northants.: one plant only on wall, Oundle, TL035880, F. H. Perring, 2001, herb. Gill Gent. 43/6.2. Salicornia ramosissima 110, Outer Hebrides: north of Lochboisdale, NF7819, E. Pilling, 2000, det. R. J. Pankhurst, E. Ist record since 1970. 43/6.6. Salicornia fragilis | *106, E. Ross: intertidal zone of muddy shore, Inver, NH8608235, B.R. & C. B. Ballinger, 2005, det. I. K. Ferguson. t44/1.1x4. Amaranthus retroflexus x A. bouchonii *29, Cambs.: several plants scattered amongst numerous plants of both parents in beet field, Cross Drove, Littleport, TL584868, A. C. Leslie, 2005, CGE. © 45/3.1. Montia fontana 32, Northants.: recently cleared area of relict heathland, Harlestone Firs, SP719639, B. J. Laney, 2004, conf. P. Stanley, herb. Gill Gent. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995). 45/3.1a. Montia fontana subsp. fontana 110, Outer Hebrides: Chuthar, South Harris, NGI9K, R. J. Pankhurst & P. A. Smith, 2004. Ist record since 1970. 45/3.1b. Montia fontana subsp. variabilis 110, Outer Hebrides: wet peaty pool near sea, Husinis, NA9IW, R. J. Pankhurst & P. A. Smith, 2004, det. A. O. Chater. Ist record since 1970. 46/1.2. Arenaria norvegica *106, E. Ross: rock outcrop, Breabag Tharsuinn, NC302190, G.P. Rothero, 1998. Still present 2005. 46/17.3. Spergularia marina *+32, Northants.: three of four large patches on road verge, Wellingborough, SP870666, S. L. M. Karley, 2004, herb. S.L.M. Karley. | £46/20.7. Silene armeria *©29, Cambs.: one plant on disturbed ground, south of the new Cutter Ferry Bridge, Midsummer Common, Cambridge, TL459590, A. C. Leslie, 2005. 46/20.10. Silene dioica 110, Outer Hebrides: Northbay area, Barra, NF70, D. Dupree, L. Gaskell, C. Velander, G. Orledge, 1998. Ist record since 1970. 746/20.11. Silene gallica: ®28, W. Norfolk: arable field edge, Snettisham, TF667335, N. Harris, 2001. £46/25.6. Dianthus barbatus *42, Brecs.: waste ground, Bwlch, SO146223, W. J. H. Price & M. Porter, 2005. +46/GYP.mur. Gypsophila muralis *70, Cumberland: dumped earth in field, Glasson Moss, NY240610, F. J. Roberts, 2005, det. G. Halliday, LANC. 47/1.12. Persicaria lapathifolia 110, Outer Hebrides: Loch an Duin, NB189408, N. F. Stewart & J. J. Day, 1995. Ist record since 1970. 47/1.16. Persicaria minor *86, Stirlings.: Shore of Loch Lomond, between Balmaha and Endrick Mouth, NS4290, A. J. Silverside, 1978. t47/3.1. Fagopyrum esculentum *©42, Brecs.: road verge, 1km E of Bronllys, SO149355, M. & C. Porter, 2005. 50, Denbs.: bare ground, Gresford, $J346536, B. Formstone, 2005. Ist record since 1930. 47/4.5. Polygonum boreale *106, E. Ross: coast, Munlochy, NH640525, J. K. Butler, 1992. 47/8.8x13. Rumex xpropinquus (R. longifolius x R. crispus) *108, W. Sutherland: sea cliff, Oldshore, NC1959, P. Merler, 1948, det. J. E. Lousley, E. Ex. herb. Nat. Con. Council. 47/8.13x19. Rumex xpratensis (R. crispus x R. obtusifolius) *108, W. Sutherland: Inchnadamph, NC250215, I. P. Green, 2005. Also found by I. P. Green at Achmelvich and Durness. 47/8.14x15. Rumex xruhmeri (R. conglomeratus x R. sanguineus) *44, Carms.: growing with both parents in disturbed dunes in vicinity of new pond near the inland control tower, RAF. Pembrey Sands, Tywyn Burrows, SN375049, BSBI meeting, 2005, det. A. O. Chater & H. J. Killick. 48/2.1. Armeria maritima +29, Cambs.: four plants scattered along the east side of the central reservation of the M11, north-west of Whittlesford, TL457486 - 457488, P. H. Oswald, 2005. 1st record since 1930 when it was seen along the tidal R. Nene at Foul Anchor. 51/1.3. Hypericum androsaemum *91, Kincardines.: wooded roadside, Strachan, NO685932, D. Welch, 2005. +51/l.for. Hypericum forrestii | *70, Cumberland: lakeshore gravel bank, Derwentwater, SE side, NY266202, F. J. Roberts, 2005, LANC. +53/1.2. Malva alcea *©44, Carms.: three plants, Shidenyn farmyard and road verge between Felin Newydd and Penygaer Farm, SN352231, C. Cheffings, M. Godfrey & A. R. Vickery, 2005. PLANT RECORDS 395 53/2.0lbxthu. Lavatera xclementii (L. olbia x L. thuringiaca) *51, Flints.: established garden throw-out on waste ground, Mold, SJ238644, J. Phillips, 1997. ©53/4.1. Aleea rosea *49, Caerns.: one plant in concrete sea-wall, near Little Orme, Penrhyn Bay, SH8281, W. McCarthy, 2005. 57/1.1. Viola odorata +106, E. Ross: damp woodland, outside wall of Contin chuchyard, NH455557, B. R. & C. B. Ballinger, 2005. 1st record since 1970. 57/1.4x5. Viola xbavarica (V. riviniana x V. reichenbachiana) *32, Northants.: many plants, well established on woodland bank, Wild Wood, Silverstone, SP672430, C. Christie, 2001, herb. Gill Gent. 57/1.4x6. Viola xintersita (V. riviniana x V. canina) *93, N. Aberdeen: grassland on fixed dunes, Sands of Forvie, NK019265, I. P. Green & D. Welch, 2005, ABD. £57/1.12x13. Viola xcontempta (V. tricolor x V. arvensis) *80, Roxburghs.: with both parents in barley stubble field, south-east of Clarilaw, Hawick, NT530180, M. E. Braithwaite, 2005, herb. R.W.M.C. *57/1/13. Viola arvensis 104, N. Ebudes (Raasay): disturbed soil, West Suisnish, NG553355, S. J. Bungard, 2004. 1st record since 1970. 61/2.11x15. Salix xlaurina (S. cinerea x S. phylicifolia) *93, N. Aberdeen: sea cliff, Whinnyfold, NK083335, D. Welch, 2003, det. R. D. Meikle, herb. D. Welch. 61/2.11x16. Salix xsubsericea (S. cinerea x S. repens) *91, Kincardines.: wooded roadside, Lochton, Banchory, NO708973, D. Welch, 2005, det. R. D. Meikle, herb. D. Welch. 61/2.12x15. Salix xludificans (S. aurita x S. phylicifolia) 69, Westmorland: edge of Skelsmergh Tarn, SD533966, N. F.Stewart & J. Lambert, 2005, det. R. D. Meikle, LANC. Ist record since 1970. 62/10.2. Matthiola sinuata *44, Carms.: one plant (in flower) on yellow dunes, Ginst Point, Laugharne, c.SN3208, J. & C. Rees, 2005. 62/12.3. Rorippa islandica *£48, Merioneth: occasional to locally frequent on rough ground beside river and concrete footpath all way down to footbridge, on E bank of Afon Glaslyn below Ynys-lleian, Beddgelert, SH54, S. P. Chambers, 2005, conf. A. O. Chater. Ist record in N. Wales. P. M. Benoit describes R. islandica as a classic example of a neophyte - a spreading weed of artificial habitats, suddenly appearing where it has never been before. +62/14.qui. Cardamine quinquefolia (Bieb.) Schmalh. *35, Mons.: one small clump on bank of R. Usk, Abergavenny, SO292139, J. S. Rees, 2005, det. T. G. Evans. @ Apparently Ist record for British Isles. 62/15.2. Arabis glabra ®28, W. Norfolk: forest ride, Foulden, TF783004, J. E. Gaffney, 2002. 62/21.4. Draba muralis *©32, Northants.: a few scattered plants on railway bank near Great Oxenden, SP737828, B. J. Laney, 2005, conf. E. J. Clement, herb. Gill Gent. £62/25.2. Capsella rubella 23, Oxon: towpath of Oxford Canal, SP5006, J. Edgington, 2005, OXF. Ist record since 1962. 62/27.1. Teesdalia nudicaulis 91, Kincardines.: dry grassland close-grazed by rabbits, Crathes, NO747962, D. Welch, 2005. Ist record since 1970. 106, E. Ross: shingle by river, Dunglass Island, NH535547, Inverness Botany Group, 2001. Ist record since 1970. 62/30.6. Lepidium latifolium ®28, W. Norfolk: roadside, Dereham, TF997123, D. & B. Mathias, 2001. 762/35.2. Sinapis alba 69, Westmorland: upland kale field, north side of Ninezergh Lane, Levens, SD495849, J. H. Clarke, 2005, LANC. Ist record since 1970. 62/41.1. Crambe maritima *93, N. Aberdeen: rocky shore, Peterhead, NK125455, M. Innes, 2005, conf. D. Welch. Very probably a recent colonisation - small plants first noticed 2003. ©62/CAL.irr. Calepina irregularis Thell. *29, Cambs.: several plants growing on discarded rootballs of trees on waste ground, Cambourne, TL313599, N. P. Millar, 2005, det. A. C. Leslie, CGE. © *65/2.2. Rhododendron luteum *80, Roxburghs.: pine woodland, The Moor, Monteviot, NT639260, J. Mercer, 2000, det. D. McKean, E, herb. R.W.M.C. Naturalised in quantity with R. ponticum. 65/13.2. Vaccinium microcarpum ®106, E. Ross: damp meadow near Alladale Lodge, NH440894, C. Sandom, B. R. & C. B. Ballinger, 2005. 396 PLANT RECORDS 69/1.3. Primula veris 106, E. Ross: riverside grassland, Brahan, NH509539, S. Cohen, 1999. Ist record since 1970. Seen in 2005 in NH66. +70/1.2. Pittosporum tenuifolium *49, Caerns.: established in wood above track to Menai Strait, Bangor, SH5873, S. P. Chambers, 2005. +71/HYD.mac. Hydrangea macrophylla *©44, Carms.: one plant in fly-tipped rubbish on vacant ground, WDA development site, Delta Lake, Machynys, SS508986, G. Hutchinson, I. K. Morgan & R. D. Pryce, 2004. 72/2.6. Ribes alpinum +29, Cambs.: one large shrub on an old earth tip beside field hedge, Oakington Road, Cottenham, TL431661, J. L. Sharman, 2004. Ist record since C19. 73/1.1. Crassula tillaea ®28, W. Norfolk: roadside, Hoe, TF988162, K. A. & G. Beckett, 2001. +73/5.3. Sedum confusum *5, S. Somerset: small patch at base of wooded slope on margin of wood, Dunster, SS989433, P. R. Green, 2005S. ©74/1.1xchixros. Astilbe xarendsii (A. japonica x A. chinensis x A. rosea) *44, Carms.: several flowering plants in disused silica quarry, Mynydd-y-Garreg, Kidwelly, SN435085, V. & A. Lewis, 2005. 74/9.1. Chrysosplenium oppositifolium 110, Outer Hebrides: Stornoway Castle, NB43, R. J. Pankhurst & W. A. J. Cunningham, 1997. Ist record since 1970. £75/3.2x3. Spiraea xbillardii (S. alba x S. douglasii) | *42, Brecs.: bank of R.Wye, Builth Wells, SO035513, M. & C. Porter, 2005. £75/6.3. Filipendula camtschatica *106, E. Ross: roadside, A835, Monadh More, NH582523, J. O. Mountfield, 2003. Still present 2005. +¢75/8.2. Rubus tricolor *42, Brecs.: beside track in open woodland, 1.5km SW Penderyn, SN935072, M. & C. Porter, 2005. 75/8.32. Rubus vigorosus *44, Carms.: edge of Molinia dominated grassland, Penygraig Goch, near Myddfai, SN743225, BSBI meeting, 2005, det. M. Porter. Occasional at this farm 75/8.60. Rubus lindleianus *91, Kincardines.: roadside, Cammachmore, NO901948, D. Welch, 2004, det. A. Newton, herb. D. Welch. 75/8.81. Rubus sciocharis *93, N. Aberdeen: abandoned railway, Maud, NJ930481, D. Welch, 2004, herb. D. Welch. 75/8.112. Rubus nemoralis *91, Kincardines.: roadside, St Cyrus, NO742649, D. Welch, 2004, conf. A. Newton, herb. D. Welch. 75/8.254. Rubus rufescens *86, Stirlings.: wood margin, Island of Clarinsh, Loch Lomond NNR, NS4189, A. McG. Stirling, 1980, det. A. Newton, E. +75/9.7. Potentilla recta *44, Carms.: one plant at base of hedge, Ty’r-waun, Pwll, Llanelli, SN470013, I. K. Morgan, 2005. *46, Cards.: streambank by level crossing, Llanbadarn Fawr, SN599807, A. O. Chater, 2005, NMW. 75/9.16. Potentilla sterilis 106, E. Ross: woodland, River Alness, NH644714, P. Wortham, 1998. Ist record since 1970. Still present 2005. t75/13.3. Geum macrophyllum *©5, S. Somerset: garden weed, not originally planted, Wellington, ST140200, S. J. Parker, 2005. May have come in with birdseed. 75/15.1. Agrimonia eupatoria 106, E. Ross: coast, Shandwick, NH856738, H. McHaffie, B. R. & C. B. Ballinger, 2005. Ist record since 1970. Also recorded for Atlas by BSBI group in NH98 in 1999. 75/19.12. Alchemilla glomerulans *70, Cumberland: herb-rich meadow at 400 m, Garrigill, NY762385, F. J. Roberts, 2005. ®106, E. Ross: cliff, Loch Luchd Coire, Seanna Braigh, NH283878, V. Halcrow, 2004. @®106, E. Ross: cliff, Coire Lochain, Ben Wyvis, NH767708, B. R. & C. B. Ballinger, 2005, herb. B.B. 75/21.4x16. Rosa arvensis x R. sherardii *5(0, Denbs.: hedge, small lane, St George, Abergele, SH967757, M. Stead, 2005, det. R. Maskew, NMW. 75/21.13bx16. Rosa caesia subsp. vosagiaca (female) x R. sherardii (male) *110, Outer Hebrides: top of low sea cliff, Glendale, NF792146, R. J. Pankhurst, 2003, det. R. Maskew, E. Ist record with subsp. of R. caesia identified. +75/21.bru. Rosa brunonii Lindl. *29, Cambs:.: at least four large plants, some climbing 30-40 ft into trees and shrubs, southern end of Coe Fen, Cambridge, TL450571, A. C. Leslie, AOS. CGE. © Apparently 1st record for British Isles. PLANT RECORDS 397 75/22.4x5. Prunus xfruticans (P. spinosa x P. domestica) *48, Merioneth: by track on edge of sand dunes, near Coleg Harlech, SH53, N. E. & R. Sherlock, 1998, conf. K. W. Page. 75/27.1. Malus sylvestris *106, E. Ross: Aultgowrie, NH45, L. Garrard, G. Hogg, S. Hogg, T. C. G. Rich, 1987. Also at east end Loch Achilty, NH43795691, 2004. +75/28.7. Sorbus intermedia s.s. 110, Outer Hebrides: Stornoway Castle, NB43, R. J. Pankhurst & W. A. J. Cunningham, 1997. Ist record since 1970. +75/28.hup. Sorbus hupehensis *95, Moray: five self-sown trees on edge of Balnacoul Wood, Mosstodloch, NJ322600, I .P. Green, 2005, det. E. J. Clement. £75/31.1. Photinia davidiana *95, Moray: twenty-four self-sown bushes in Balnacoul Wood, Mosstodloch, NJ3259, I. P. Green, 2005. +75/32.10. Cotoneaster salicifolius *7, N. Wilts.: bird-sown on wall-top, Marlborough, SU186689, J. E. Oliver, 2003. £75/32.16. Cotoneaster integrifolius 7, N. Wilts.: bird-sown on stonework, Coate Water, Swindon, SU177826, J. E. Oliver, 2003. Ist record since 1970. *91, Kincardines.: dyke by stream, Maryculter, NJ894009, D. Welch, 2005. +75/32.17. Cotoneaster linearifolius *7, N. Wilts.: bird-sown seedling on stonework, Coate Water, Swindon, SU177826, J. E. Oliver, 2003. Parents cultivated nearby. [£75/32.25. Cotoneaster nanshan 77, Lanarks.: error in VCCC. No known record. Delete. ] £75/32.34. Cotoneaster bullatus *91, Kincardines.: riverside woodland, Banchory, NO710957, D. Welch, 2005. Self-seeded from nearby landscaping. +75/32.40. Cotoneaster sternianus *50, Denbs.: quarry, Llandulas, SH912777, W. McCarthy, 2003. [+75/32.ind. Cotoneaster induratus 77, Lanarks.: error in VCCC. No known record. Delete. | [£75/32.lid. Cotoneaster lidjiangensis 77, Lanarks.: error in VCCC. No known record. Delete. | £77/3.1xori. Colutea xmedia (C. arborescens x C. orientalis) *5, S. Somerset: flowering, at edge of car park, Porlock Weir, SS864478, C. J. Giddens, 2005. 77/10.3. Ornithopus perpusillus 106, E. Ross: by forest track, West Kinkardine Hill, NH588872, B. R. & C. B. Ballinger, 2005. Ist record since 1970. 77/14.13. Vicia lutea 91, Kincardines.: embankment of abandoned railway, St Cyrus, NO737635, B. Hogarth, 2005. Ist record since 1970. ©77/14.fab. Vicia faba *7, N. Wilts.: spoil, Bradford-on-Avon, ST823632, J. Presland, 2003. *49, Caerns.: disturbed ground near new superstore, Porthmadog, SH5638, Caerns. Recording Group, 2005. +77/15.9. Lathyrus latifolius *106, E. Ross: dune by sea, Shandwick, NH859750, B. R. & C. B. Ballinger, 2004. Confirms M. J. Crawley Atlas record for NH87 in 1999. 77/16.3. Ononis spinosa *t44, Carms.: sparsely vegetated opencast spoil, Ffos-las former opencast site, Trimsaran, SN453062, I. K. Morgan, 1999. ?Introduced in seed mix. 77/16.4. Ononis repens 106, E. Ross: urban site, Muir of Ord, NH5348, P. Wortham, 1995. Ist record since 1970. Still present, Muir of Ord station, 2005. 77/18.3. Medicago minima @®29, Cambs.: many fine plants on a dry, open, SW facing bank above pond, Cambridge Science Park, Milton, TL464617, A. C. Leslie, 2005, CGE. 77/19.8. Trifolium fragiferum 50, Denbs.: grassland, Pentre Mawr Parc, Abergele, SH943786, J. A. Green, 2005. Ist record since 1970. £77/21.1x2. Lupinus xregalis (L. arboreus x L. polyphyllus) *106, E. Ross: by shore, Barbaraville, NH749721, B. R. & C. B. Ballinger, 2005. Garden escape. Confirms M. J. Crawley Atlas record for NH77 in 1999. 77/25.4. Genista anglica 106, E. Ross: Hillockhead, NH747603, Inverness Botany Group, 1997. Ist record since 1970. Still present 2005. £77/25.ten. Genista tenera (Jacq.) Kuntze *17, Surrey: one large shrub and several smaller _ ones self-seeded from nearby gardens, Turf Hill, Bagshot, SU932617, G. Hounsome, 2005, det. E. J. Clement. @ 77/26.3. Ulex minor 32, Northants.: recently cleared area of relict heathland, Harlestone Firs, SP718639, G. M. Gent, 1999. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995). +77/PHA.vul. Phaseolus vulgaris *42, Brecs.: canal verge, Brecon, SO047282, A. & V. Lewis, 2005. 398 PLANT RECORDS ©77/PIS.sat. Pisum sativum *49, Caerns.: disturbed ground near new superstore, Porthmadog, SH5638, Caerns. Recording Group, 2005. +78/1.1. Hippophae rhamnoides —__ 110, Outer Hebrides: Berneray, NF909813, R. J. & A. F. Pankhurst, 2004. Ist record since 1970. 781/1.2. Lythrum hyssopifolium ©®29, Cambs:.: in gravel scrape by Elney Lake, Fen Drayton gravel pits, TL336693, S. Brown, 2005, det. N. P. Millar. Considered native in v.c.29. 81/1.3. Lythrum portula 32, Northants.: recently cleared area of relict heathland, Harlestone Firs, SP718689, G. M. Gent, 2005, herb. Gill Gent. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995). 81/1.3b. Lythrum portula subsp. longidentatum *110, Outer Hebrides: ditches west side of North Uist, NF76, W. A. Shoolbred, 1898, NMW, det. D. E. Allen (as var. longidentata), in Allen Watsonia 3: 85-91 (1954). +83/2.gun. Eucalyptus gunnii *5, S. Somerset: self-sown on side of passageway as street weed, Wellington, ST135195, S. J. Parker, 2005. 84/1.2. Epilobium parviflorum 106, E. Ross: dunes, lochan south-east of control tower, Morrich More, NH8482, BSBI group, 1999. Ist record since 1970. [84/1.2x6. Epilobium xdacicum (E. parviflorum x E. obscurum) *52, Anglesey: damp ground by track, Ty Mawr, Paradwys, SH416708, I. R. Bonner, 2004, herb. I.R.B. In Watsonia 26: 82 (2006). Delete - plant re-determined as Epilobium xfloridulum (E. parviflorum x E. ciliatum) by G. D. Kitchener. ] 84/1.2x7. Epilobium xpersinicum (E. parviflorum x E. roseum) *29, Cambs.: several plants with both parents at edge of pavement bordering the Cherry Hinton Brook, Burnside, Cambridge, TL475574, A. C. Leslie & C. D. Preston, 2005, CGE. 84/1.2x8. Epilobium xfloridulum (E. parviflorum x E. ciliatum) *44, Carms.: working limestone quarry, Cilyrychen Quarry, Llandybie, SN616166, A. O. Chater, K. A. Cottingham & R. D. Pryce, 2001, det. A.O. Chater. *52, Anglesey: damp ground by track, Ty Mawr, Paradwys, SH416708, I. R. Bonner, 2004, herb. I.R.B. det. G. D. Kitchener. Originally determined as Epilobium xdacicum (E. parviflorum x E. obscurum). 84/1.3x6. Epilobium xaggregatum (E.montanum x E. obscurum) *44, Carms.: one plant at woodland edge, Penygraig Goch, near Myddfai, SN743225, BSBI meeting, 2005, det. A. O. Chater. 84/1.7. Epilobium roseum *50, Denbs.: roadside, Pwllglas, SJ126554, J. A. Green, 2005. 84/1.7x8. Epilobium xnutantiflorum (E. roseum x E. ciliatum) *29, Cambs.: one plant with both parents at edge of pavement bordering the Cherry Hinton Brook, Burnside, Cambridge, TL475574, A. C. Leslie & C. D. Preston, 2005, CGE. +84/1.8. Epilobium ciliatum 104, N. Ebudes (Raasay): waste ground, Clachan, NG549364, S. J. Bungard, 2002, det. T. D. Pennington. Ist record for Raasay. Also known as garden weed, West Suisnish. 84/1.8x9. Epilobium xfossicola (E. ciliatum x E. palustre) *44, Carms.: rushy area in field, Cwrtbrynybeirdd, near Trapp, SN665178, BSBI meeting, 2005, det. A. O. Chater. t84/5.1. Fuchsia magellanica *7(), Cumberland: two bushes on bank of R. Cocker, Cockermouth, NY 122306, D. Broughton, 2005. +85/1.3. Cornus alba *42, Brecs.: roadside scrub, Bronllys, SO151345, A. & V. Lewis, 2005. 85/1.5. Cornus suecica *70, Cumberland: 10 - 15 sterile shoots on ledge in gully at 800m on Great End, NY227085, S. Webb, 2005, LANC. t91/2.3. Euphorbia corallioides *29, Cambs.: numerous plants naturalised both on and beside grass track along field margin at the western end of village, Barton, TL402560, A. C. Leslie, 2005, CGE. +91/2.0bl. Euphorbia oblongata *7, N. Wilts.: urban structures, Malmesbury, ST936871, J. E. Oliver & J. Wall, 2004. Declining here. Known for several years. 94/2.1. Radiola linoides 91, Kincardines.: wet floor of abandoned quarry, Maryculter, NO880993, D. Welch, 2005. Ist record since 1970. 95/1.1. Polygala vulgaris 106, E. Ross: coast, Shandwick, NH856738, C. Dixon, J. W. McIntosh, G. P. Rothero, 1998. Ist record since 1970. Still present 2005. +99/1.mac. Acer macrophyllum *8, S. Wilts.: numbers of seedlings from two planted trees, Rushmore Estate, ST958194, J. E. Oliver, 2003. PLANT RECORDS 399 +100/1.1. Rhus typhina 47, Monts.: one young stem coming up through soil-mound, farm sales outlet S of Welshpool, SJ222056, S. P. Chambers, 2005. ©101/1.1. Ailanthus altissima *7, N. Wilts.: seedlings on riverside and roadside, Chippenham, ST920735, M. Waters, 2003. +102/1.8. Oxalis articulata 50, Denbs.: garden escape, St George, SH973758, J. A. Green, 2005. *102/1.14. Oxalis incarnata *49, Caerns.: woodland edge, Marl Woods, SH8078, NWWT group, 2005. +103/1.8. Geranium himalayense *5, S. Somerset: single clump on edge of track to garages, Taunton, ST235245, S. J. Leach, 2003. 103/1.9. Geranium sanguineum ©7,N. Wilts.: roadside, Kingsdown, ST817671, J. Presland, 2003. Ist record since 1970. +103/1.13xpla. Geranium xmagnificum (G. ibericum x G. platypetalum) 106, E. Ross: by track, Strathgarve Lodge. NH404612, M. Dean, B. R. & C. B. Ballinger, 2005, herb. B.B. Ist record since 1970. ©103/2.tri. Erodium triflorum Cav. *29, Cambs.: two plants self sown at base of fence in front of 320 Coldham’s Lane (and several more at base of hedge in front of number 314), Cambridge, TL474580, A. C. Leslie, 2005. Naturalised. © ©104/1.maj. Tropaeolum majus *£110, Outer Hebrides: by shore, Brevig, Barra, NL694985, R. J. Pankhurst, 2004. +106/1.1. Hedera colchica *42, Brecs.: road bank, 2km S of Llangattock, SO204166, S. P. Chambers, 2005. +107/1.ran. Hydrocotyle ranunculoides *29, Cambs.: Soham Lode, first seen 2003 and by 2004 was dominating the mill pond and extending downstream to railway line, Soham, TL589730 - 585730, R. Torrens, 2003. *32, Northants.: very well established in large pond, Higham Ferrers A45 Road, SP950687, Environment Agency, 2003, herb. Gill Gent. *49. Caerns:.: organic farm pond and surrounding land, Llangybi, SH4340, D. Thorpe, 2005. 107/5.3. Chaerophyllum temulum 106, E. Ross: roadside, west end of Loch Achilty, NH435571, J. Muscott, 1996. Ist record since 1970. Still present here 2005. 107/15.1. Sium latifolium ®28, W. Norfolk: dike by pasture, West Harling, TL967853, Norfolk Flora Group & R. W. Ellis, 2002. 107/19.5. Oenanthe crocata 32, Northants.: well established and spreading on shaded river bank, Bluebell Walk, Peterborough, TL1999, D. Broughton, 2005, conf. B. Partridge. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995). 107/20.1. Aethusa cynapium +106, E. Ross: by track near shore, Tain, NH785824, B. R. & C. B. Ballinger, 2005, herb. B.B. Ist record since 1970. 107/31.1. Cicuta virosa ®28, W. Norfolk: waterside, Cockley Cley, TF787037, J. E. Gaffney, 2004. 107/39.2. Peucedanum palustre ®28, W. Norfolk: pool in pasture, Oxburgh Hythe, TF732000, Norfolk Flora Group & K. A. & G. Beckett, 2004. 107/40.1. Pastinaca sativa +106, E. Ross: waste ground by shore, Inver, NH864828, B. R. & C. B. Ballinger, 2005, herb. B.B. Ist record since 1970. 107/44.1a. Daucus carota subsp. carota 110, Outer Hebrides: Uig sands, NB050328, 1971. lst record since 1970. 108/3.3. Centaurium littorale 106, E. Ross: widespread on short turf at top side of saltmarsh, Morrich More, NH846842, S. Smyth, B. R. & C. B. Ballinger, 2005, herb. B.B. Ist record since 1970. ©111/IPO.pur. Ipomoea purpurea *5, S. Somerset: single self-sown plant in pavement crack, Dunster, SS989435, P. R. Green, 2005. £113/2.1. Nymphoides peltata *91, Kincardines.: fishery pool created c. 1998, Raemoir, - Banchory, NO696977, D. Welch, 2005. 114/1.1. Polemonium caeruleum *+110, Outer Hebrides: rough ground near football pitch, Stornoway, NB432334, E. Pratt, 1999. £116/4.2. Symphytum asperum *95, Moray: one plant growing on bank of the River Spey, under road bridge of the A941, Craigellachie, NJ286452, I. P. Green, 2005. 400 PLANT RECORDS 116/4.3. Symphytum tuberosum *+110, Outer Hebrides: waste ground by sports centre, Coll, NB463386, R. J. Pankhurst, 2003, E. Previously reported from Scarastavore, South Harris by Heslop Harrison ef al. in Journal of Botany 79:164-169 (1941), but without a voucher. This is the first confirmed record. +116/10.1. Trachystemon orientalis | *69, Westmorland: copse by road junction, Slackhead, Beetham, SD493787, J. H. Clarke, 2005. 116/15.9. Myosotis ramosissima 106, E. Ross: coast, North Sutor, Nigg, NH806688, P. Wortham, 1997. Ist record since 1970. Still present 2005. +116/16.1. Omphalodes verna 70, Cumberland: near fisherman’s hut, R. Eden, Coombs Wood, Armathwaite, NY506446, G. Lines, 2005. Ist record since 1970. ©117/1.hyb. Verbena xhybrida *5, S. Somerset: single self-sown plant in pavement crack, Dunster, SS990435, P. R. Green, 2005. ©117/CAR.inc x mon. Caryopteris xclandonensis (C. incana x C. mongholica) A. Simmonds ex Rehder *29, Cambs.: one young flowering plant, self sown on low brick wall in front of Sherlock Court, Cambridge, TL436599, A. C. Leslie, 2005, CGE. @ £118/1.2. Stachys byzantina *49, Caerns.: on dumped soil in field, Pydew, SH8179, W. McCarthy, 2005. 7118/2.1a. Ballota nigra subsp. meridionalis 44, Carms.: at edge of disused, vegetating tarmac car-park, Pembrey Burrows Local Nature Reserve, SS414993, BSBI AGM field meeting, — 2005. Ist record since 1970. +118/6.2. Galeopsis angustifolia ®8, S. Wilts.: on edge of arable field margin, farmland, DTE Salisbury Plain, SU247494, J. Moon, 2004. 118/11.3. Teucrium scordium *52, Anglesey: pool margin, Bryngwran, SH349768, R. Birch, 2005. Ist Welsh record. Status unknown - ?native, as probably introduced by wildfowl. 118/12.2. Ajuga pyramidalis @®108, W. Sutherland: hazel scrub, River Laxford, NC254457, P. A. & I. M. Evans, 2005. Other plants in the vicinity. +118/23.3. Mentha spicata 110, Outer Hebrides: East Pabbay, NL68, R. J. Pankhurst & J. Swindells, 1998. Ist record since 1970. +118/23.3x4. Mentha xvillosa (M. spicata x M. suaveolens) 110, Outer Hebrides: Castlebay, Barra, NL6698, R. J. Pankhurst, 1998. Ist record since 1970. 120/1.pal. Callitriche palustris *86, Stirlings.: sandy shore of Endrick Mouth, Crom Mhin Bay, Loch Lomond NNR, NS426901, J. Mitchell, 2005, det. R. Lansdown, E, NMW. 2nd record for British Isles. | 121/1.1. Plantago coronopus 32, Northants.: well established on verge of A6, Corby, SP8888, R. D. Cleaver, 1999, conf. G. M. Gent, herb. Gill Gent. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995). 106, E. Ross: coast, Nigg ferry, NH805689, P. Wortham, 1997. Ist record since 1970. Still present 2005. 121/1.3b. Plantago major subsp. intermedia *95, Moray: frequent on tracks in Lossie Forest, NJ280677, I. P. Green, 2005. *110, Outer Hebrides: Ceann Iar, Monach Islands NNR, NF60526200, P. A. Smith, 2005, det. T. C. G. Rich, NMW. Cult. T. C. G. Rich. £122/1.3. Buddleja globosa *©7, N. Wilts.: hedgerow, NW of Bromham, ST983679, J. Presland, 2003. *51, Flints.: naturalised, Leete path, 1.6km SW of Rhydymwyn, SJ195657, G. Wynne, 2003. +124/1.1. Verbascum blattaria 35, Mons.: one plant on the edge of a pile of stony earth, N of Millbrook Garden Centre and Trothy Brook, SO501112, T. G. Evans & C. Titcombe, 2005. Ist record since 1970. +124/1.10. Verbascum speciosum *t42, Brecs.: road verge, Brecon, SO033284, A. & V. Lewis, 2004. 124/1.11. Verbascum pulverulentum ®28, W. Norfolk: disused sand pit, Blackborough End, TF680143, Norfolk Flora Group & R. M. Leaney, 2005. £124/1.pho. Verbascum phoeniceum *69, Westmorland: overgrown dump of soil, Ninezergh Lane, Levens, SD494849, G. Halliday, 2005, LANC. 124/2.1. Scrophularia nodosa 110, Outer Hebrides: foot of wall, Willowglen Road, Stornoway, NB434340, R. J. Pankhurst, 2001, E. Ist record since 1970. +124/4.2. Mimulus guttatus *29, Cambs.: margin of pond, Cambridge Science Park, Milton, TL466617, D. Barden, 2004. Probably deliberately introduced but now well established. PLANT RECORDS 401 124/5.1. Limosella aquatica *69, Westmorland: shallow muddy pool, Tarn Syke, Sunbiggin, NY667076, G. Quartly-Bishop, 2005, LANC. ©124/13.mar. Linaria maroccana *44, Carms.: waste ground, Burry Port Harbour, SN443003, V. & A. Lewis, 2005. With Centaurea cyanus, Viola tricolor subsp. tricolor, Sedum album and S. rupestre. +124/16.18. Veronica peregrina 29, Cambs.: one plant as a weed on Monksilver Nursery, Oakington Road, Cottenham, TL436663, J. L. Sharman, 2005, CGE. 3rd v.c. record and Ist since 1958. +124/16.24. Veronica hederifolia 106, E. Ross: coast, Portmahomack/Rockfield, NH98, BSBI group, 1999. Ist record since 1970. +124/17.elixspec. Hebe xfranciscana (H. elliptica x H. speciosa) *29, Cambs.: one self sown plant at base of wall, Coveney, TL487820, A. C. Leslie, 2005. 124/20.5x1. Euphrasia arctica x E. rostkoviana *46, Cards.: horse-grazed pasture with bracken at 240 m, below road 1km SE of Devil’s Bridge, SN747763, A. O. Chater, 2003, det. A. J. Silverside, NMW. 124/20.5bx19. Euphrasia xvenusta (E. arctica subsp. borealis x E. scottica) *46, Cards.: flush on S-facing grassy slope at 370m, 200m W of Nant-llwyd, Soar y Mynydd, SN781524, A. O. Chater, 2003, det. A. J. Silverside, NMW. 124/20.5x20. Euphrasia arctica x E. heslop-harrisonii *110, Outer Hebrides: above salt marsh, Seilebost, NGO82965, A. F. Pankhurst, 2005, det. A. J. Silverside, E. 124/20.7xla. Euphrasia nemorosa x E. rostkoviana subsp. rostkoviana *46, Cards.: grassy working area in conifer forest at 430m, Bryn Du-bach, 3km SSE of Llanddewi-Brefi, SN678527, A. O. Chater, 2003, det. A. J. Silverside, NMW. 124/20.7x8. Euphrasia nemorosa x E. pseudokerneri *46, Cards.: gravelly verge of FC road by lead mine at 270m, with both parents growing nearby, Mynach Vale, 3km E of Devil’s Bridge, SN772775, A. O. Chater, 2003, det. A. J. Silverside, NMW. 124/20.7x9. Euphrasia nemorosa x E. confusa *79, Selkirks.: locally common in basic pasture at 540m, Red Scar, Hopetown Craig, Ettrick, NT185071, R. W. M. Corner, 2005, det. A.J. Silverside, herb. R.W.M.C. 110, Outer Hebrides: short turf, Hellisay, Sound of Barra, NF764036, P. A. Smith, 2003, det. A. J. Silverside, E.1st record since 1970. 124/20.8. Euphrasia pseudokerneri ®28, W. Norfolk: chalk grassland, Cockthorpe Common, TF984430, Norfolk Flora Group & R. W. Ellis, 2000. ®28, W. Norfolk: chalk grassland, Grimes Graves, TL817899, S. R. J. Woodell, 2000. 124/20.11. Euphrasia frigida 110, Outer Hebrides: at 450m on stony hilltop, Caiteshal, NB240046, R. J. Pankhurst & P. A. Smith, 2002. Ist record since 1970. 124/20.12. Euphrasia foulaensis 106, E. Ross: top end of saltmarsh, Morrich More, NH8483, P. A. Evans, 1998. Ist record since 1970. Still present and abundant 2005 (B. R. & C. B. Ballinger), confirmed A. J. Silverside. | @110, Outer Hebrides: Liniclate, Benbecula, NF7849, E Pilling, 2000, det. A. J. Silverside, E. 124/20.12x14. Euphrasia foulaensis xE. ostenfeldii *110, Outer Hebrides: turf at cliff edge, Scotasay, NG182927, R. J. Pankhurst, 2005, det. A. J. Silverside. 124/20.19. Euphrasia scottica 106, E. Ross: damp woodland, Loch Luichart, NH385580, B. Goater, S. Erskine, M. Godfrey, 1999. 1st record since 1970. ©124/21.1. Odontites jaubertianus *32, Northants.: short unimproved calcareous grassland on edge of disused WW? airfield, Spanhoe, SP940978, R. Mileto, 2005, conf. E. J. Clement, herb. E.J.C. 124/23.1. Parentucellia viscosa *29, Cambs.: two plants on disturbed ground near Fen Causeway, Coe Fen, Cambridge, TL448574, J. Shanklin, 2005. 124/25.2. Pedicularis sylvatica 32, Northants.: wet field, Yardley Chase MOD area (SSSI), SP836556, M. Coleman, 1997. Rediscovered having been presumed extinct (Flora of - Northamptonshire, 1995) and Ist record since Druce 1930. +£124/DIA.int. Diascia integerrima E. Mey. ex Benth. *@©42, Brecs.: waste ground, Penderyn, SN939087, M. & C. Porter, 2005. Ist Welsh record. @ Apparently Ist record for British Isles. £124/SUT.cor. Sutera cordata Kuntze *©42, Brecs.: river bank, 3km W of Hay-on-Wye, SO198408, M. & C. Porter, 2005, det. G. M. Kay. @ +125/1.2. Lathraea clandestina *110, Outer Hebrides: Askernish, South Uist, NF738238, W. Neill, 2005. Probably introduced with Salix trees. 402 PLANT RECORDS +127B/CAT.big. Catalpa bignonioides Walter *29, Cambs.: two self sown young plants at base of wall in car park behind shops, Norfolk Street, Cambridge, TL461584, A. C. Leslie, 2005. 128/2.6. Utricularia minor 106, E. Ross: Corriehallie Forest, NH396488, P. Wortham, 1999. Ist record since 1970. Still present in this locality 2004. ¢129/1.3. Campanula lactiflora *91, Kincardines.: shingle by R. Dee, Kingcausie Island, Maryculter, NJ867012, D. Welch, 2005, ABD. ©129/9.1. Downingia elegans *32, Northants.: two or three plants found in recently seeded area in marshy field, Weston Favell Mill, SP792608, R. Moore, 1997, det. M. Mullin, herb. Gill Gent. 130/3.1. Sherardia arvensis 91, Kincardines.: unsprayed turnip field, Crathes, NO750960, D. Welch, 2005, ABD. Ist record since 1970. 130/7.1. Cruciata laevipes +106, E. Ross: coast, Kiltearn, NH615652, P. Wortham, 1997. Ist record since 1970. Still present 2005. t131/1.1. Sambucus racemosa *48, Merioneth: edge of plantation, beside the Bala — Llangynog road near Llandderfel, SH981352, D. O. Elias, 2005. Known here for 2 or 3 years. +131/3.orbxmic. Symphoricarpos xchenaultii (S. orbiculatus x S. microphyllus) 27/5 INI, Wilts.: two clumps on old canal towpath, Swindon, SU136835, J. E. Oliver, 2004. t131/5.1. Leycesteria formosa *91, Kincardines.: shore by policies, Johnshaven, NO804675, | D. Welch, 1997. +131/6.1. Lonicera pileata *7, N. Wilts.: one self-seeded young plant on stonework, Chippenham, ST920735, J. E. Oliver & G. Yerrington, 2003. *69, Westmorland: wall-top, Kendal, SD517930, C. E. Wild, 2005, LANC. *+131/6.4. Lonicera xylosteum *£80, Roxburghs.: old hedge, east of Clarilaw, Hawick, NT530180, M. E. Braithwaite, 1990. Status unchanged in 2006. *133/1.2. Valerianella carianata *32, Northants.: waste ground, Sibbertoft, SP6882, F. Pearson, 1998, conf. G. M. Gent, herb. Gill Gent. £134/2.1. Cephalaria gigantea *8, S. Wilts.: seeding heads on plants on derelict site, Jockey Green, SU285639, J. E. Oliver, 2004. ¢135/1.3. Echinops bannaticus *44, Carms.: on SW facing sandy bank, Pembrey Country Park, SN406010, I. K. Morgan, 1998, det. G. Hutchinson, NMW. Still present in 2005. The original record was published in error as E. sphaerocephalus (Welsh Bulletin 76: 33 (2005)). A throw-out or deliberate planting? | *©46, Cards.: one plant in alleyway, self-sown from nearby garden, N of High Street, Lampeter, SN576481, C. M. Overton, 2005, det. A. O. Chater, NMW. 135/5.3. Carduus crispus 93, N. Aberdeen: arable grassland, Rattray, NK087581, I. P. Green, 2005. Ist record since 1970. *135/11.3. Centaurea cyanus *©110, Outer Hebrides: bank opposite Cuddy Point, Stornoway Castle, NB418328, R. J. Pankhurst, 2005, E. Apparently introduced with seed mixture. +135/17.1. Picris echioides *©104, N. Ebudes (Skye): single plant on roadside, S of Sconser, NG534293, S. J. Bungard, 2004. Site later tarred. +135/22.1. Lactuca serriola 42, Brecs.: industrial estate road verge, Brynmawr, SO1991 14, M. & C. Porter, 2005. Ist record since 1970. *49, Caerns.: roadside bank, Penrhyn Bay, SH8181, M. Stead, 2005. *52, Anglesey: roadside gravel, Valley, SH286789, R. Birch, 2005. Second plant at SH289786. 135/25.58. Taraxacum cambricum *69, Westmorland: meadow by canal, Stainton, SD523853, A. M. Boucher, 2005, det. A. J. Richards, herb. A.M.B. 135/25.100. Taraxacum spiculatum *69, Westmorland: pavement weed, Natland, SD521891, A. M. Boucher, 2005, conf. A. J. Richards, LANC, herb. A.M.B. +135/25.104. Taraxacum acutifidum *69, Westmorland: grassland, Rinkfield Road, Kendal, SD519912, A. M. Boucher, 2005, det. A. J. Richards, LANC, herb. A.M.B. 135/25.120. Taraxacum cophocentrum *69, Westmorland: footpath border, Natland, SD519890, A. M. Boucher, 2005, conf. A. J. Richards, herb. A.M.B. £135/25.122. Taraxacum corynodes *69, Westmorland: verge near church, Natland, SD521892, A. M. Boucher, 2005, det. A. J. Richards, herb. A.M.B. *135/25.127. Taraxacum densilobum *69, Westmorland: coastal grassland, Milnthorpe, SD488815, A. M. Boucher, 2005, det. A. J. Richards, herb. A.M.B. *135/25.138. Taraxacum adiantifrons "*86, Stirlings.: waste ground by old lemonade factory, Milngavie, NS552748, R. J. Pankhurst, 2003, det. A. J. Richards, E, herb. R.J.P. PLANT RECORDS 403 £135/25.149. Taraxacum laeticolor *69, Westmorland: roadside verge, B6385, Crooklands, SD533826, A. M. Boucher, 2005, det. A. J. Richards, herb. A.M.B. 135/25.155. Taraxacum leucopodum *44, Carms.: moderately fertile and well drained grazing/hay field with some winter grazing and no record of recent ploughing, Waunygwiail Fach Farm, Four Roads, SN447086, A. Stevens, 2004, det. A. J. Richards, NMW. Good material of this scarce plant. 135/25.172. Taraxacum oblongatum *+110, Outer Hebrides: Harris Hotel, Tarbert, NB152000, P. A. Smith, 2005, det. A. J. Richards, E. 135/25.217. Taraxacum undulatum *69, Westmorland: coastal grassland, Milnthorpe, SD483823, A. M. Boucher, 2005, det. A. J. Richards, LANC, herb. A.M.B. *135/25.coa. Taraxacum coartatum *110, Outer Hebrides: Harris Hotel, Tarbert, NB152000, P. A. Smith, 2005, det. A. J. Richards, E. +135/27.6b. Pilosella aurantiaca subsp. carpathicola *108, W. Sutherland: rough ground between path and backs of gardens, Lochinver, NC095230, I. M. Evans, 2005. 135/28.29. Hieracium placerophylloides *46, Cards.: ungrazed rocky river bank at 285m, Afon Pysgotwr Fawr, 300m SE of Bryn-glas, SN738511, A. O. Chater, 1995, det. D. J. McCosh, NMW. 135/28.111. Hieracium pollinarioides *110, Outer Hebrides: Glen Skeaudale, NB140034, R. J. Pankhurst & P. A. Smith, 1996, det. D. J. McCosh, E. *135/28.115. Hieracium scotostictum *46, Cards.: roadside banks, Derwen-fach, Llandygwydd, SN242436, A. O. Chater & P. A. Smith, 1998, det. D. J. McCosh, NMW. Escaped from cultivation in adjacent garden. 135/28.167. Hieracium schmidtii 110, Outer Hebrides: Beinn na h’ Aire, North Uist, NF901590, P. A. Smith, 1997, det. D. J. McCosh, E. Ist record since 1970. £135/28.asp. Hieracium asperatum Jord. ex Boreau *46, Cards.: ballast of disused railway in cutting, behind Penyrangor, Aberystwyth, SN581808, A. O. Chater, 1993, det. D. J. McCosh, NMW. Previously det. as H. pollichiae by P. D. Sell in 1994. Site now destroyed. © 135/28.avi. Hieracium aviicola Jord. ex Boreau *46, Cards.: roadside hedgebank at 275m, 600m WSW of chapel, Trefenter, SN601685, A. O. Chater, 1998, det. D. J. McCosh, NMW. @ 135/28.con. Hieracium consociatum Jord. ex Boreau *46, Cards.: ballast of disused railway, Aberbachnog, 2km E of Llandysul, SN373403, A. O. Chater, 1985, det. D. J. McCosh, NUW. © 135/28.deg. Hieracium deganwyense Pugsley *46, Cards.: mortared walls of wheelpit of ruined lead mine at 370m, Nantycreuau mine, 4km E of Ponterwyd, SN789802, A. O. Chater, 1991, det. D. J. McCosh, NMW. Previously det. as H. scoticum by P. D. Sell in 1994. @ 135/28.rio. Hieracium rionii Gremli *29, Cambs.: one plant by side of School Lane, Histon, TL441630, P. D. Sell, 2004, CGE. @ £135/29.1. Gazania rigens *©49, Caerns.: pavement edge near beach café, Penrhyn Bay, SH8281, W. McCarthy, 2005. 135/30.4. Filago minima = 32, Northants.: recently cleared area of relict heathland, Harlestone Firs, SP718638, B. J. Laney, 2004, conf. P. Stanley, herb. Gill Gent. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995). 135/33.2. Gnaphalium sylvaticum 7, N. Wilts.: woodland path, Cobham Frith, SU255669, J. Newton, 2003. Ist record since 1970. ©135/34.ita. Helichrysum italicum *29, Cambs.: one self sown plant at base of wall, Front Court, Emmanuel College, Cambridge, TL452583, A. C. Leslie, 2005. [+135/35.1. Inula helenium *91, Kincardines.: alder wood on island in R. Dee, Maryculter, NJ866009, D. Welch, 1995. Misidentification of Telekia speciosa (Watsonia 21: 397 (1997)). Delete from VCCC. | £135/38.1.Telekia speciosa *91, Kincardines.: alder woodland on island in R. Dee, Kingcausie Island, Maryculter, NJ866009, D. Welch, 2005, ABD. Original 1995 record called Inula helenium in error. *93, N. Aberdeen: streamside in gorge, Craig, NJ477245, D. Welch, 2005, herb. D. Welch. The original 1987 record for this site (the 2° record for v.c. 93) was erroneously called Inula helenium (Watsonia 17: 478 (1989)). £135/44.1 Conyza canadensis *93, N. Aberdeen: flower bed, Invervrie, NJ775216, I. P. Green, 2005. *£135/50.2. Artemisia verlotiorum *32, Northants.: many plants, well established on road verge, Middleton Cheney, SP509413, C. Christie, 2001, conf. G. Hansen, herb. Gill Gent. 404 PLANT RECORDS 7135/55.2. Anthemis arvensis *©110, Outer Hebrides: bank opposite Cuddy Point, Stornoway Castle, NB418328, R. J. Pankhurst, 2005. Apparently introduced with seed mixture. *135/55.4.Anthemis tinctoria *©110, Outer Hebrides: path by west harbour, Tarbert, NB148002, R. J. Pankhurst, 2005. ©135/55.aus. Anthemis austriaca Jacq. *29, Cambs.: several plants on old soil heap, Pembroke College sports field, Grantchester Road, Cambridge, TL438570, A. C. Leslie, 2005, CGE. @ +135/62.3. Senecio inaequidens *©32, Northants.: drained pond, Woodford Halse, SP505505, M. Atkins, 2003, conf. B. Partridge, herb. Gill Gent. *44, Carms.: about 3 large plants in disturbed ground at east end of Burry Port Southern Distributor Road construction site, SN453004, I. K. Morgan, 2005, NMW. *@©46, Cards.: eight plants on waste ground, Ynys-las boatbuilding yard, SN616932, A. O. Chater, 2005, NMW. *50, Denbs.: garage forecourt, Denbigh, SJ054669, J. A. Green, 2005. ¢135/67.1. Sinacalia tangutica *29, Cambs.: apparently self sown at base of wall in narrow, concreted basement area beside Emmanuel College, Parker Street, Cambridge, TL454583, A. C. Heshes 2003: +£135/68.1. Ligularia dentata *69, Westmorland: south-west margin of pond, Witherslack, SD434862, J. Adams & W. Nelson, 2005. £135/71.2. Petasites japonicus *106, E. Ross: damp ground by river, Evanton, NH617654, | B.R. & C. B. Ballinger, 2005, herb. B.B. £135/71.3. Petasites albus 110, Outer Hebrides: Stornoway Castle, NB43, R. J. Pankhurst & W. A. J. Cunningham, 1997. Ist record since 1970. +135/74.1. Ambrosia artemisiifolia *108, W. Sutherland: four plants in garden near bird feeders, Craggan Glas, Clashnessie, NCO53308, E. King, 2005, det. P. A. & I. M. Evans, herb. I.M. Evans. ©135/76.2. Xanthium spinosum **32, Northants.: garden, Hunsbury, Northampton, SP727579, B. J. Laney, 2005, conf. E. J. Clement, herb. Gill Gent. ¢135/78.1. Rudbeckia hirta *95, Moray: one plant only on rubbish tip, Elgin, NJ236631, I. P. Green, 2005. +135/79.2. Helianthus tuberosus *44, Carms.: one large plant, c.2m across, in derelict ground used as council ash tip in 1960s, near Pemberton, Llanelli, SS530999, I. K. Morgan, 2005, NMW. 142/1.1x2. Potamogeton xgessnacensis (P. natans x P. polygonifolius) | *69, Westmorland: in 3m of water, Little Langdale Tarn, NY309033, C. Newbold, 1999, det. C. D. Preston, LANC. Ist English and 3rd British record. 142/1.7. Potamogeton alpinus 32, Northants.: pond, Yardley Chase MOD area (SSSI), SP839559, M. Wade, 1996. Rediscovered having been presumed extinct (Flora of Northamptonshire, 1995) and 1st record since Druce 1930. 142/1.17. Potamogeton compressus 29, Cambs.: one plant immediately west of the sluice on north side of Moreton’s Leam, Stanground Sluice, just E of Peterborough, TL209974, P. Kirby, 2004. Ist record since 1912. 146/1.1. Zostera marina ®106, E. Ross: intertidal zone, Portmahomack, NH915846, B. R. & C. B. Ballinger, J. O. & S. A. Mason, 2005, herb. B.B. +147/2.1. Lysichiton americanus *93, N. Aberdeen: ditch in policies, Pitcaple Castle, NJ726262, D. Welch, 2005. Escape from garden. £147/5.2b. Arum italicum subsp. italicum *106, E. Ross: woodland edge, Rosemarkie, NH734578, B. R. & C. B. Ballinger, 2005, herb. B.B. Garden escape. £147/7.1. Arisarum proboscideum +42, Brecs.: shaded rough ground, Gurnos, SN778094, A. & V. Lewis, 2005. ©147/PIS.str. Pistia stratiotes L. *4, N. Devon: two small rosettes, Grand Western Canal, SS9812, J. Ison, 2005. © 151/1.6. Juncus foliosus 104, N. Ebudes (Raasay): mud by burn, Arnish, NG596482, S. J. Bungard, 2003. 1st record for Raasay and Ist for v.c.104 since 1970. 151/1.8. Juncus ambiguus 104, N. Ebudes (Rona): shore, Big Harbour, NG616565, S. J. Bungard, 2003. 1st record for Rona and Ist for v.c.104 since 1970. 104, N. Ebudes (Raasay): shore, Oskaig, NG547377, S. J. Bungard, 2003. Ist record for Raasay. PLANT RECORDS 405 151/1.13x14. Juncus xsurrejanus (J. articulatus x J. acutiflorus) *44, Carms.: frequent in rushy field (with both parents), Cwrtbrynybeirdd near Trapp, SN665180, BSBI meeting, 2005, det. A. O. Chater. 151/1.20. Juncus maritimus *104, N. Ebudes (Skye): tidal river, Camasunary, NG510191, A. M. B. Averis & A. M. Averis, 1996. Seen by S. J. Bungard 2006. 152/4.1. Bolboschoenus maritimus *#32, Northants.: old fishing lake, Heyford, SP657624, T. Balbi, 1997, conf. G. M. Gent. £152/11.1. Cyperus longus *7(), Cumberland: one large clump on S bank of R. Derwent, Cockermouth, NY 120308, D. Broughton, 2005. +152/11.2. Cyperus eragrostis *29, Cambs.: self sown seedlings from plants presumably deliberately introduced, Lake Lambert, Cambourne, TL319598, N. P. Millar & A. C. Leslie, 2005. 152/14.1. Cladium mariscus 108, W. Sutherland: patch 20 x 20m in shallow pool, saddle mire, Mullach na Creige Deirge, NC165473, P. A. & I. M. Evans, 2005. Ist record since 1970. 152/16.32. Carex sylvatica 106, E. Ross: Craig Wood, Dingwall, NH556600, Dundee Naturalists Society, 2005. Ist record since 1970. Previous Atlas record, M. J. Crawley in NHSS, ooo) 152/16.57. Carex limosa 106, E. Ross: bog, Scardroy Burn, NH180533, P. Wortham, 2003. Ist record since 1970. Still present 2005. 152/16.60. Carex atrata *86, Stirlings.: Ben Lomond, NN30, L. Watt, 1888, Noted by A. McG. Stirling in Scottish Naturalist IV. 152/16.65x67. Carex xelytroides (C. acuta x C. nigra) *69, Westmorland: with both parents in marsh west of Roman camp, Ambleside, NY371034, M. S. Porter, 2005, conf. A. O. Chater, LANC. 152/16.65x68. Carex xprolixa (C. acuta x C. elata) *52, Anglesey: fen, Cors Castell, SH4980, R. H. Roberts, 1982?, det. A. C. Jermy, NMW. From an un-named herbarium sheet passed on by Mrs Roberts. 152/16.67x69. Carex xdecolorans (C. nigra x C. bigelowii) 50, Denbs.: grassy summit, Cadair Bronwen, SJ0O78346, S. Stille, 2005, det. A. O. Chater, M. S. Porter & R. W. M. Corner, NMW. Ist Welsh record. 153/12.2x3. Festuca xfleischeri (F. arundinacea x F. gigantea) *70, Cumberland: at least 2 plants on sandy W bank of R. Eden, Wetheral, NY467549, F. J. Roberts, 2005, det. A. Copping & T. A. Cope, LANC. One of three sites. 153/12.7a. Festuca rubra subsp. rubra 110, Outer Hebrides: Allt Volagir, South Uist, NF/99295, R. J. Pankhurst, 2000, det. C. A. Stace, E. Ist record since 1970. 153/12.8. Festuca ovina 106, E. Ross: dunes, Morrich More, NH88, BSBI group, 1998. Ist record since 1970. Widespread here 2005. 153/12x14/7x1. xFestulpia hubbardii (Festuca rubra x Vulpia fasciculata) *95, Moray: growing on sand dunes with both parents, Findhorn, NJO37649, I. P. Green, 2005, conf. A. Copping. 7153/14.3. Vulpia myuros *©110, Outer Hebrides: car park, Tarbert, NB150002, R. J. Pankhurst, 2005, det. C. A. Stace, E. 153/14.4b. Vulpia ciliata subsp. ambigua *32, Northants.: two or three large patches in open rough ground, Polebrook Airfield, TL099871, R. D. Cleaver, 1997, det. C. A. Stace, herb. Gill Gent. £153/18.10. Poa palustris *95, Moray: one plant on edge of small pond in Lossie Forest, NJ296665, I. P. Green, 2005, conf. T. A. Cope. ©153/25.cil. Melica ciliata var. atropurpurea L. *44, Carms.: several plants, self-sown from cultivated plants nearby, in cobbled/gravelled farmyard, Glynhir Mansion, Llandybie, SN639152, BSBI meeting, 2004, det. G. M. Kay. @ 153/26.1. Helictotrichon pubescens 106, E. Ross: dunes, Morrich More, NH852835, BSBI group, 1998. Ist record since 1970. Still present 2005. 153/26.2. Helictotrichon pratense. 106, E. Ross: coast, Shandwick, NH856738, C. Dixon, J. W. McIntosh, G. P. Rothero, 1998. Ist record since 1970. Still present here 2005. +153/44.1. Apera spica-venti *©32, Northants.: found during survey of farmland, Sacrewell Farm, TL080998, T. C. E. Wells, 1999, herb. T.C.E. Wells. *©44, Carms.: one plant only in disturbed, sparsely vegetated ground, Dyfatty, Burry Port, SN460096, BSBI meeting, 2005, det. G. M. Kay, NMW. 406 PLANT RECORDS +153/50.7. Bromus secalinus 28, W. Norfolk: arable field, Bradenham, TF916105, K. A. & G. Beckett, 2002. Ist record since 1970. 153/56.1x3. Elytrigia xlaxa (E. repens x E. juncea) 106, E. Ross: sandy shore, Nigg near ferry terminal. NH796687, B. R. & C. B. Ballinger, 2005, herb. B.B. 1st record since 1970. 153/58.1. Hordelymus europaeus 32, Northants.: recently cleared area in woodland, large area of over 100 plants, Short Wood, TLO191, A. Colston, 1998, conf. F. H. Perring, herb. Gill Gent. Ist record since 1970. £153/62.ric. Cortaderia richardii *3, S. Somerset: single self-sown clump on side of woodland ride, Pendomer, ST521089, P. R. Green, 2005, det. I. P. Green. 153/67.2. Spartina anglica *£106, E. Ross: muddy shore, Dingwall, NH557590, B. R. & C. B. Ballinger, 2004, det. T. A. Cope, herb. B.B. Previously recorded as Spartina alterniflora var. glabra. +153/68.1.Echinochloa crus-galli *©31, Flints.: reclaimed industrial site, Rhydymwyn, SJ205668, G. Wynne, 2003. ¢153/70.pum. Setaria pumila ©51, Flints.: reclaimed industrial site, Rhydymwyn, SJ205668, G. Wynne, 2003. Ist record since 1917. +153/70.ver. Setaria verticillata *69, Westmorland: pavement weed, Finkle St, Kendal, SD515926, A. M. Boucher, 2005, LANC, herb. A.M.B. +153/71.san. Digitaria sanguinalis *69, Westmorland: pavement weed, Finkle St, Kendal, SD515926, A. M. Boucher, 2005, LANC, herb. A.M.B. £153/FAR.mul. Fargesia murielae (Gamble) T. P. Yi *44, Carms:.: originally planted, now naturalised and spreading on bank clothed in tall herb and bramble to west of larger of two ponds, Penarth Ganol, Pumsaint, SN649401, K. A. & R. D. Pryce, 2005, det. G. Hutchinson, NMW. Ist record for naturalised plant away from coastal amenity plantings. @ 154/1.1c. Sparganium erectum subsp. neglectum 50, Denbs.: riverside, Llandulas, SH909787, J. A. Green, 2005. Ist record since 1911. 154/1.2. Sparganium emersum 106, E. Ross: small loch, Black Loch, NH792756, Loch Survey, 1994. Ist record since 1970. Still present 2005. +158/10.1. Tulipa sylvestris *93, N. Aberdeen: roadside, Pitcaple, NJ734254, I. P. Green, 2005. £158/12.1. Lilium martagon *93, N. Aberdeen: roadside, Bourtie, NJ783244, I. P. Green, 2005. £158/18.2. Ornithogalum angustifolium *110, Outer Hebrides: graveyard, Eriskay, NF784116, R. J. & A. F. Pankhurst, 2004. +158/19.1. Scilla bifolia © *©110, Outer Hebrides: rough grassland near Ormiclate, South Uist, NF73, D. L. Smith, 1956, GLAM. 158/19.4. Scilla verna 44, Carms.: two plants, one with c.25 and the other with 2 flowering stems, on crest of cliff in limestone grassland, Dolwen Point, Pendine, SN233078, I. K. Morgan & R. N. Stringer, 2005, NMW. Ist record since E. B. G. Jones’ c.1950 record from Llanstephan. £158/22.1. Chionodoxa forbesii *69, Westmorland: by copse at road junction, Slackhead, Beetham, $D493787, J. H. Clarke, 2005. £158/24.5. Allium subhirsutum *44. Carms.: three clumps associated with A. vineale in disturbed ground at edge of platform, Ferryside railway station, SN366104, G. Hutchinson, 2005, NMW. 2nd Welsh record 158/24.11. Allium oleraceum 91, Kincardines.: steep deciduous woodland, Morphie, NO717620, D. Welch, 2005, ABD. Ist record since 1970. ©158/24.cep. Allium cepa *£42, Brecs.: shingle along R.Usk, near Scethrog, SO106244, M. & C. Porter, 2005. ©158/24.tub. Allium tuberosum Rottl. Ex Spreng *29, Cambs.: several plants, apparently self-sown on steps of 45 Panton Street, Cambridge, TL453574, P. H. Oswald, 2005, det. A. C. Leslie. © £159/2.5. Sisyrinchium striatum *44, Carms.: established on colliery tip, south of Y Garth, near Ammanford, SN619133, I. K. Morgan, 2005, det. G. Hutchinson, NWW. *159/5.4. Iris versicolor 110, Outer Hebrides: shore, Benbecula, NF847553, S. J. Bungard, 1998. 1st record since 1970. PLANT RECORDS 407 £159/13.1. Crocosmia paniculata *93, N. Aberdeen: trackside, Strichen, NJ957563, D. Welch, 2005. 162/3.7. Epipactis phyllanthes ®28, W. Norfolk: fen edge, Dereham, Lolly Moor, TF992103, D. & B. Mathias, 2002. 162/11.1. Corallorhiza trifida ®106, E. Ross: vegetated dunes, in 4 locations under birch and willow, Morrich More, NH852834, B. R. & C. B. Ballinger, 2005. 162/15x18/1x2. xPseudorhiza bruniana (Pseudorchis albida x Dactylorhiza maculata) *104, N. Ebudes (Skye): coastal grassland, Ullinish, NG319368, B. Allen, 1994. 162/17x18/1x1. xDactyloglossum mixtum (Coeloglossum viride x Dactylorhiza fuchsii) 110, Outer Hebrides: Luskentyre, NGO72987, R. S. C. Ferreira, 1990. Ist record since 1970. 162/18.1x2. Dactylorhiza xtransiens (D. fuchsii x D. maculata) 110, Outer Hebrides: Fuday, NF7308, R. Waterston, 1980, E. Ist record since 1970. 162/18.1x5. Dactylorhiza xvenusta (D. fuchsii x D. purpurella) 110, Outer Hebrides: North Uist, NF845766, S. J. Bungard, 1998. Ist record since 1970. 162/18.3b. Dactylorhiza incarnata subsp. coccinea 104, N. Ebudes (Raasay): moorland, South Fearns, NG5835, S. J. Bungard, 1997, det. R. M. Bateman. Ist record since 1970. 162/18.5x6. Dactylorhiza purpurella x D. majalis 110, Outer Hebrides: Clachan, Trumisgarry, North Uist, NF867749, R. J. Pankhurst, 1979. Ist record since 1970. 162/18.7. Dactylorhiza traunsteineri ®28, W. Norfolk: part alkaline mire, Roydon, TF686227, R. A. Yaxley, 2000. ®28, W. Norfolk: alkaline mire, Castle Acre, TF806152, H. E. Williamson, 2000. Watsonia 26: 409-415 (2007) 409 Book Reviews Change in the British Flora 1987-2004 (A report on the BSBI Local Change survey). M. E. Braithwaite, R. W. Ellis & C. D. Preston. Pp. iv + 382. Botanical Society of the British Isles, London. 2006. £12. ISBN 0-901158-34-8. Botanists have always been fascinated by the distribution of species, as witnessed by the Herculean efforts of B.S.B.I. members in gathering records for the two Atlases, and the plethora of county Floras complete with lists of localities and dot maps at tetrad or 1-km square resolution. Although we might get weary of it at times, square-bashing remains something of a national pastime. Make no mistake; the end-products of all this recording activity (the Red Data Books, Atlases and County Rare Plant Registers) are abso- lutely vital to those of us working to conserve the British flora. But while dot maps tell us a great deal about the distribution of each species, they say rather less about how that distribution is changing through time, and next to nothing about the environmental factors ‘driving’ those changes. Desirable as it may be, the scope for developing a botanical equivalent to the Breeding Bird Survey or Butterfly Monitoring Scheme is extremely limited. Counting birds and butterflies is much easier, on the whole, than counting plants. Many plants are simply ‘uncountable’ — without digging them up you can’t tell where one individual ends and the next begins, and taking account of the seed- bank is practically impossible. Multiply these problems by the sheer numbers of species involved and you begin to see why botanists have generally opted to record presence/ absence within sample plots and grid-squares to generate distribution and frequency data — rather than attempting direct counts to give abundance data of the sort favoured by zoologists. But there is a great deal that can be gleaned from dot-map frequency data, as demonstrated by the New Atlas, which included a quantitative analysis of changes in the British flora between the periods 1930-69 and 1987-99, along with a ‘change index’ for those taxa mapped in the 1962 Atlas and again in the New Atlas. The New Atlas analysis was a real eye-opener, with species like Chrysanthemum — segetum, Gnaphalium sylvaticum, Ranunculus arvensis, Pedicularis sylvatica, Scleranthus annuus and Spergula arvensis being listed in the ‘top 100’ species showing the greatest relative decrease between the two recording periods. This paved the way for a root-and-branch revision of the Red Data List for Great Britain in 2005. The analyses of change in the New Atlas and the Red List were based on 10-km square data; yet there has always been a suspicion that data at this scale could be masking (or under- estimating) some of the changes taking place — after all, within a 10-km square a common species occurring in 25 tetrads could, in theory, be lost from all but one tetrad (a colossal decline) without having any effect on either its 10-km dot distribution or its frequency at 10- km scale. This is a problem, of course, with dots at any scale, and indicates the need for a variety of surveys at different spatial scales. It also suggests that declines in more widespread species may be more readily discernible by examining presence/absence data from smaller sampling units such as tetrads or 1-km squares. A difficulty with smaller units, of course, is that blanket coverage of the country becomes impossible, so one therefore has to adopt some kind of sampling approach — which is where the B.S.B.I.’s “Local Change’ project, and the volume under review, makes its entrance onto the stage. On the face of it, “Local Change’ has moved us to another level: the change indices are derived from tetrad data, and the time interval between the first (1987-88) and second (2003- 04) surveys is just 18 years, covering the most recent date-class in the New Atlas (1987-99) and the five years following. This is a colourful and attractively produced book, well written, with lots of colour photographs and clearly set- out tables; but it is a technical report, and beneath its apparent accessibility there lurks a degree of complexity that many readers will find hard to penetrate. The data analyses, in particular, are bewildering, and even those with a reasonable grasp of mathematics are likely to struggle to make sense of the statistical methods (and underlying assumptions) employed, although — to be fair to the authors — these are fully explained in an Appendix. Great care has been taken to point out the limitations of both the data and the analytical methods 410 BOOK REVIEWS being used. Throughout the report there are caveats and ‘cautionary notes’, and readers need to bear these in mind when interpreting and using the results. In particular, for any measure of % change there is also a measure of the confidence that can be attached to it, and all too often the 90% confidence limits point to considerable uncertainty as to the reliability and extent of the changes being reported. Species are grouped according to Broad Habitats, then divided into subgroups by means of a TWINSPAN analysis of their British distributions as given in the New Atlas. Grouping species in this way has certain advantages, of course, in that one can then look at changes across a whole suite of species having similar national distributions and habitat requirements; but, to be honest, I found the TWINSPAN groups really rather unhelpful, and would have preferred a simpler analysis (as in the New Atlas), with species examined individually first, and then en bloc to discern those ecological, phytogeographical, morph- ological (etc.) attributes particularly associated with ‘stable’, ‘decreasing’ or ‘increasing’ species. You can, of course, pick your way through the report in search of the ecologically (and statistically) significant — and tabulated results for all species, in alphabetical order, are helpfully given in an Appendix — but at times it is a bit like searching haystacks for needles. Nevertheless, there is much in this report to quicken the pulse, and much, too, of relevance not just to conservationists ‘on the ground’ but also to those working in policy areas like climate change and sustainable land-use. For example, as hinted at in the New Atlas, it seems clear that quite a few species are now increasing markedly in response to climate change, or at any rate the recent run of warmer temperatures, including ‘southern’ or Medi- terranean plants like Geranium rotundifolium, Medicago arabica, Oenanthe pimpinelloides, Ranunculus parviflorus, Ophrys apifera and Anacamptis pyramidalis. Many increasing neophyte taxa have their origins in warm- temperate or subtropical regions, e.g. Campanula poscharskyana, Conyza sumatrensis, Allium triquetrum and Geranium pyrenaicum, and it is likely that these, too, are benefiting from the milder weather. Declining species are predominantly those associated with nutrient-poor (infertile) habitats such as calcareous grasslands and_ heather moorland: species like Alchemilla xantho- chlora, Briza media, Centaurea _ scabiosa, Cirsium acaule, Pedicularis _ sylvatica, Rhinanthus minor and Viola lutea. This agrees with the general findings in the New Atlas but, interestingly, the ‘Local Change’ survey has thrown up a number of species which, while ‘stable’ or showing only slight decline at 10- km square scale, have apparently undergone significant recent declines at tetrad scale. Some species of arable/horticultural land continue to decline, including Chrysanthemum segetum, Galeopsis speciosa and Silene noctiflora, but others are apparently increasing, possibly benefiting from agri-environment schemes introduced since the original survey (e.g. set- aside, arable field margins, etc.). Many declining species, as one would expect, seem to have been mostly lost from tetrads away from their core areas, usually in parts of the country where they were already pretty thin on the ground. Are ‘nature conservation’ activities making a difference to these species? Certainly, it would be interesting to know whether ‘edge-of-range’/outlying populations are becoming increasingly rest- ricted to ‘protected sites’ — SSSIs, County Wildlife Sites, Protected Road Verges, etc. — or agri-environment land. That many rare/scarce plants are now found mainly on protected sites has been well demonstrated in some areas (see, for example, the entries in the Dorset Rare Plants Register), but for more widespread ‘decliners’ the relationship has yet to be investigated — a worthwhile subject for future study, perhaps? The ‘Local Change’ survey came hard on the heels of ‘Atlas 2000’, and to many B.S.B.I. foot soldiers there seemed to be little time for recuperation between the end of one campaign and the start of the next. But the effort has surely been worth it. The authors do their best to tease out the main conclusions of the survey, while being honest and up-front about its limitations. The results seem to raise as many questions as they answer, but there is much here to inform and provoke, and anyone interested in Britain’s flora and how it is changing should get hold of a copy. One thing is clear: change happens simultaneously at a range of spatial scales, and if we are to document and understand it we need to have a whole batch of projects working at different scales and employing different recording methods and analytical techniques — of which the ‘Atlas’ is one, and ‘Local Change’ is another. No rest for the square-bashers, then? S. J. LEACH BOOK REVIEWS Al] The Botanist. The botanical diary of Eleanor Vachell (1579-1948). Edited by M. Forty & T. Rich. Pp. 227. National Museum of Wales. 2006. Softback. £19.50. ISBN 0-7200-0565-5. Eleanor Vachell’s diary has long been known to the happy few from a typescript in the National Museum of Wales, and now, thanks to the diligence of Michelle Forty and Tim Rich, this incomparable document of some 200,000 words is available to the wider botanical public. I am not aware of any other writing that so vividly communicates the enjoyment, the excitements and the detailed mechanics of plant-hunting in Britain and Ireland, except perhaps parts of the generally rather more sober New Naturalist book Mountain Flowers by John Raven and Max Walters. Eleanor Vachell was the daughter of a distinguished Cardiff physician and amateur botanist with whom she started, at the age of twelve, compiling what was to become an outstanding private herb- arium. She began a diary at the same time. The work under review is not so much daily entries as a Series of narratives written up in the third person, mostly apparently after the event. Her driving forces were a passionate desire to see every British plant, an ambition to colour in her illustrated “Bentham” (in fact Fitch’s //lust- rations of the British Flora, and Butcher & Strudwick’s Further Illustrations), and above all her friendship and good-natured rivalry with other botanists. She was a serious investigator, corresponding with and sending specimens to experts on many difficult groups, including Potamogeton, Rosa, Fumaria, Centaurium, Rhinanthus, Callitriche and Limosella. The diary covers most of the country, and Vachell’s most regular companions on the innumerable, often successful, often disastrous, but always high-spirited expeditions were four other lady enthusiasts who _ constituted themselves as the Trudgeleers (a term of expr- essive but obscure Irish origin). Others who figure prominently include Francis Druce, T. J. Foggitt and his wife, A. J. Wilmott, N. D. Simpson and R. W. Butcher, and many of the other botanists of the period, from the aged J. E. Griffith to the young Paul Richards, appear at some time or other. Various Botanical Exchange Club events, the International Botan- ical Congress at Cambridge and a BSBI meeting in Merioneth are vividly described. G. C. Druce makes many appear-ances, none more memorable than on a wet and unsuccessful outing to see Potentilla rupestris on the Wye when he behaved with singularly poor grace. Vachell was short of funds until an inheritance from an uncle in 1938, but by 1927 she had saved up enough from writing regular botanical notes for the Western Mail (which she did from 1921 to 1948) and from wireless talks to buy a third-hand Clyno car. She changed to a second- hand Swift Coupé in 1932, and then to a Morris later the same year. As an account of the perils and joys of early motoring alone the book is worth reading. Predictably, hunts for familiar rarities such as Cypripedium (one of the few plants she never saw in spite of the most extreme efforts) and Trichomanes (at Killarney the party was expensively tricked into being taken on horseback to see planted-out specimens by a noisy group of predatory locals including a trumpeter and a _ photographer) figure prominently. But more interesting and less fashionable taxa such as the splits of Thal- ictrum minus and the upland Deschampsia species get equal attention. Repeated expeditions to most of the chief localities for rarities, such as Lochnager, which she climbed seven times, and Ben Lawers, are described in great detail. It is a measure of the sheer gusto of the writing that the 1921 account of a catastrophic climb up Lawers in appalling weather, when no plants were seen and the author was reduced at one point to crawling up to the summit cairn in thick mist carrying her vasculum in her teeth, is even more vicariously enjoyable than the account of the successful expedition two days later when virtually all the rarities were seen. The circumstantial detail of sites and populations makes many of the entries of real historical interest. The editors have provided an index that includes genera, personal names and, especially helpfully, the main counties visited. The personal names understandably lead one sometimes to just a name rather than a part- icular person, for example “Mr Taylor” refers to Peter as well as Sir George Taylor. A few interesting names are missing, such as E. H. T. Bible of Aberdyfi on p.125, and Robert Dick, deceased but touchingly apostrophised on p.107. In the text “The botanical nomenclature has largely been updated to modern genera, though sometimes where the historical context might be lost ... the original names have been retained”, but it is not indicated where this updating, which to some extent applies to species as well as genera, has or has not been done; it would have been more helpful, and editorially correct, to have retained the original names and given the modern ones in brackets. A. O. CHATER 412 BOOK REVIEWS A Natural History of Nettles. Keith G. R. Wheeler. Pp. x + 300 plus CD-ROM with 143 colour plates. Trafford Publishing, Victoria, Canada. 2004. Softback. ISBN 1-4120-2694-6. This is a monograph on the genus Urtica and its relatives on an epic scale. The author says that he has ‘tried to produce a book for naturalists [and] scientists ... with a large number of photographs and illustrations [that] would appeal to the general public.’ Thus, it combines the enthusiasm and _ polymath expansiveness of a Victorian natural historian with the latest techniques in microscopy and scientific enquiry. The author has explored an enormously diverse range of topics, often introducing a topic in one chapter and returning to it in more detail later. This is not a book to read straight through, therefore, but, rather, one to browse and dip into for sections of particular interest. It will, perhaps, be of special interest to teachers who will find a great wealth of fascinating facts and ideas for students of all ages. On the botanical side, the author starts with the classification of the Urticales and descrip- tions of the floral structures and pollination mechanisms in Urtica, with wonderful illustrations. This is followed by a brief account of Urtica dioica, U. urens and U. pilulifera in Europe with an interesting section on var. dodartii; this was once thought to be a different species but it does not breed true and breeding experiments in the early 20th century showed that the pellitory-like leaf characters were controlled by a single recessive gene. Roman nettle is thought to be extinct now in Britain, apart, perhaps, from botanic gardens: in the 1970s I saw a small colony that was maintained in a corner of Pembroke College garden, Cambridge. Chapters 14 and 15 cover the worldwide and British distribution of U. dioica, its nutritional requirements, estab- lishment, growth and general ecology. The book concludes with two chapters on the ‘sex life’ and breeding systems in Urtica spp., and an appendix on the other British ‘nettleworts’, Parietaria judaica and Soleirolia_soleirolii, again with superb illustrations. Three whole chapters are devoted to the micromorphology and functioning of stinging trichomes, and the composition and effects of the toxins in the Urticaceae and in other, quite unrelated, families. The microscopic details of trichomes are illustrated in six full page photographs and drawings but the colour illustrations on the CD-ROM must be viewed to appreciate them fully. As a child in south India, I was shown and warned against ‘elephant nettle’, Girardinia diversifolia, and having seen it again while trekking through rainforest in Tamil Nadu in 2000, I was particularly fascinated to read the account of ‘tropical tormentors’ in chapter 7. The New Zealand nettle tree, Urtica ferox, which I have also seen, is clearly not in the same league for size and fierce stinging properties as the giant stinging trees, Dendrocnide spp., of Australia. Horses can even die from contact with these plants. On the other hand, many invertebrates are unaffected by the stinging hairs. The description of the way in which a caterpillar overcomes the defences of Cnidoscolus urens (Euphorbiaceae) is another fascinating story. Other chapters describe the place of nettles in folklore and literature and the use of nettles for food and fibre. Nettle fibre is stronger than flax fibre and its use from early historical times is well known but its production in Germany during the First World War may not be so well known. In 1915, apparently, ten million acres in Austria were thought to be ‘splendidly adapted to the cultivation of the nettle’, twice that needed to replace cotton imports, but 2.7 million kg were harvested without any special cultivation. Chapter 11 describes the life history, distri- bution and field studies on associated species such as Greater Dodder, Cuscuta europaea and fungal rusts. This is followed by two chapters on the insects associated with Urtica, again with several full page plates, which can only be fully appreciated in colour on the CD-ROM. The painting of the leaf hopper Eupteryx aurata is especially beautiful. In this field again the author contributes interesting first hand accounts of territorial and courtship behaviour among vanessid butterflies and unusual oviposition on nettles by Painted lady butt-erflies, Cynthia cardui, during a particularly large invasion in 1996. There is a very large reference list and a good subject index. Critical readers may, however, be rather surprised by the inconsistent use of italics for scientific names. This seems almost random except in the case of Urtica and relatives, which are generally printed in Roman font. There are also quite a lot of typographical errors, some of which have been corrected by hand. Among _ authors’ names quoted in the text, Perrin should be BOOK REVIEWS 413 Perring, for example, and Rackman should be Rackham. Since this book is available through ‘on-demand publishing’, it may be possible to correct some of these minor faults in future productions. The author says that this book ‘has been much maligned by the conventional book publishers’. It does, indeed, fall outside the norm in today’s economy-conscious and specialist publishing world but I hope others will, like me, obtain ‘pleasure from entering the world of the nettle patch.’ B.N. K. DAVIS Atlas Florae Europaeae. Distribution of Vascular Plants in Europe. Volume 13. Rosaceae (Spiraea to Fragaria, excl. Rubus). Edited by A. Kurtto, R. Lampinen & L. Junikka. Pp. 320. The Committee for Mapping the Flora of Europe and Societas Biologica Fennica Vanamo, Helsinki. 2004. Softback. 120 euros. ISBN 951-9108-14-9. Atlas Florae Europaeae issued its first volume in 1972 and reached the end of the first volume of Flora Europaea with volume 12 in 1999. There has now been a further long pause, occasioned partly by the death of Jaakko Jalas and the retirement of Juha Suominen, who were the founding editors. The commencement of maps _ covering volume 2 of Flora Europaea has allowed a new look and small modifications to the layout. These include an amended grid system (principally for the purpose of fitting in with atlases of other groups and largely affecting areas of sea) and very useful coverage map of all the taxa covered in this part — which allows one to see straight away likely lacunae — together with the customary list of deviations from Flora Europaea. The territory divisions and the acronyms tie up with the Euro+Med PlantBase, newly set up in 2000. The maps themselves have, at last, totals of the dots under the various date and status categories. This volume is the first of four for the Rosaceae, a family made deeply complicated by apomixis; indeed two of the next three parts are to be devoted solely to Alchemilla and Rubus. That will leave Sorbus, Cotoneaster and the Pyrus/Malus complex for the last volume. This first part has 286 maps of which 54 are of Rosa and 150 of Potentilla. All other genera are small. The coverage of most genera is introduced by a general map for all the taxa covered, which allows one to see both the richness of that genus in each country and how well each has covered it - particularly relevant for critical groups. After Spiraea and Filipendula comes Rosa, and for readers from Britain and Ireland, a treatment very difficult to get to grips with, particularly under Section Canina. The treat- ment here broadly follows Flora Europaea and thus fundamentally differs from that in Stace’s New Flora and Graham & Primavesi’s B.S.B.I. Handbook. To add to our complications, the general map that prefaces the account shows that France, our nearest neighbour, has little enthusiasm for the genus other than a very broad treatment. Up to the Canina group there are no problems, other than the interesting point that the widely naturalised R. rugosa is barely recorded for France, which must be just record- ing bias. After the Canina group the synon- ymy is more or less comparable with ours and there is little to comment on other than perhaps the non-recording of R. sherardii from France. I do not think this review is the place (even assuming I had the skill) to provide a detailed comparison for the two treatments of the Canina group. Suffice it to say that each species or aggregate has a detailed (I was tempted to say bewildering) section on taxonomy, through which the skilled or patient could trace the treatment of the species concepts more familiar to us. Thus their R. dumalis is our R. caesia subsp. glauca and their R. subcanina is our R. caesia subsp. glauca x canina (R. xXdumalis sensu. Graham & Primavesi pro parte). Next come _ smaller genera _ including Sanguisorba and -Geum. Nothing particular to say there, but they are followed by the large Potentilla genus prefaced by P. fruticosa, P. palustris and P. rupestris split off into Dasiphora, Comarum and Drymocallis respectively. These alterations date from 1990 but do not seem to have been even discussed here. Potentilla argentea is split into P. argentea and P. neglecta, with the latter reaching W & S France and Scandinavia but not (yet) here. The native range of our alien species P. norvegica, P. intermedia and P. recta is well shown. P. tabernaemontani 1s mapped as our Spring Cinquefoil and P. neumanniana is shown only as a local apomict in Germany and Scandinavia. Interestingly, P. anglica is shown in France as very rare with many extinctions. 414 Perhaps one has to be grateful for this magnificent work of scholarship — in many ways much more full and satisfactory than the original written Flora — and ignore the fact that on current progress it might be complete by 2120. Personally I find the current progress deeply unsatisfying. I would much prefer a statement now of as many species as can be easily dealt with, leaving the critical groups as aggregates if necessary. Though there have been other distribution atlases for Northern Europe and Scandinavia it is a real bore that these do not cover around 40% of our flora, BOOK REVIEWS and even if they do, they are often based on data that are very old. Finally, the price. I am sure that it can be justified for the scientific work involved and the limited market. But it is totally out of court for anybody with other than the deepest pocket, especially as British natural history booksellers cite prices between £110 and £145, and this does not help dissemination of the valuable distribution and taxonomic work contained in this Atlas. D. A. PEARMAN The Wild Flower Key, revised and expanded edition. F. Rose & C. O’Reilly. Pp. 576. Frederick Warne. 2006. Softback. £19.99. ISBN 0-7232-5175-0. Since its publication in 1981 The Wild Flower Key has become one of the most popular field guides to the British flora. By combining taxonomic keys with illustrations of diagnostic features, ‘Rose’, as it is affectionately known, has helped a generation of British botanists to develop the critical skills required for field identification. This new edition incorporates much new information on the distribution, conservation and taxonomy of British species including many aliens which are now well established in the wild in the UK. Much of the work for the new edition has been carried out by Clare O’Reilly (née Coleman), a freelance botanist who abandoned a career in environmental law to complete it. Clare has edited and expanded many of the species accounts and, with the help of leading botanists, improved the keys to difficult genera. In addition, there are new chapters on ‘Plant Status’, “Ancient Woodland Indicators’, ‘Wild Plants and the Law’ and ‘Finding and Recording Wild Plants’ as well as useful lists of botanical references and websites for beginners. Although the introductory keys remain unaltered, the nomenclature and family orders have been revised to follow Stace (1997), only diverging where the authors disagree (e.g. Gymnadenia) or where recent taxonomic changes have been large (e.g. Anacamptis). This is a great improvement on the rather inconsistent taxonomic treatment of the first edition. However, most of the diagnostic characters are still based on CTW (second edition) and are therefore woefully out of date. This is a shame given the wealth of new information recently made available. The keys to families, genera and species were one of the best features of the first edition and these have remained largely unaltered (although see Primavesi 2006). More controversial will be the removal of north-west European species not present in the UK, such as Arnica montana and Digitalis lutea, although surprisingly two British neophytes have also been removed (Holosteum umbellatum, Legousia speculum- veneris). The line drawings of common native grasses have also been dropped, presumably because the wonderful companion volume to grasses is now available (Rose 1989). These deletions have made way for over 50 aliens which have become widely naturalised over the last fifty years (e.g. Buddleja davidii, Fallopia Japonica, Hirschfeldia incana, Hyacinthoides x massartiana, Persicaria wallichii, Solidago canadensis) and illustrations of 20 native montane species that were omitted from the first edition (e.g. Betula nana, Diapensia lapponica, Koenigia islandica, Linnaea borealis, Loiseleuria procumbens, Moneses uniflora, Oxyria digyna, Silene acaulis).. The coverage of the new edition is therefore less biased to the south and more reflective of the British flora at the start of the twenty-first century, including all natives and _ long- established introductions, all non-natives in over 300 10-km squares and non-natives in 50- 299 10-km squares if they are invasive or easily confused with natives. Substantial changes to the layout have made the book much easier to use in the field. The illustrations are more spaced out on the page and some poor quality pictures have been reproduced more accurately (e.g. Viola spp.) although one or two have suffered from enlargement (e.g. Myrica gale, Thesium humifusum). Virtually all the line drawings BOOK REVIEWS 415 have been redrawn and are now much clearer (see Fumaria spp.) and the ‘ID Tip’ boxes which provide handy hints and _ illustrated glossaries to diagnostic features for around twenty large families will no doubt prove popular with beginners. The design is refreshingly modern whilst retaining much of the original format and, if you treat your books as badly as I do, you’ll appreciate the plastic cover which appears to come as standard. Some of the deficiencies of the first edition remain although most are only minor draw- backs. The tone of the new introduction is rather immodest and I’m sure less energetic colleagues will resent the increase in size (it is considerably heavier than Clive Stace’s (1999) Field Flora, for example). Unfortunately there are still many typographic errors, some of which were not corrected by the publisher in order to meet the publication deadline (these are listed in Coleman (2006)). Many illustrations are on separate pages to the accompanying text, which can prove irritating, and some users may find the plethora of abbreviations (e.g. in describing distribution, habitats, status etc.) a little bewildering. The selection of illustrated species remains frustratingly incomplete and many taxa will Smlljebem too poorly covered for more experienced botanists, in particular infra- specific and critical taxa (e.g. Alchemilla spp., Erophila spp., Montia fontana). One wonders whether these could have been better covered at the expense of very rare species such as Orchis laxiflora or the vegetative keys to flora of specific habitats that, although being one of the best features of the original, are probably rarely used. Also a simple table of ecological attributes (e.g. soil pH, moisture, fertility, life- form, etc.) would have been more informative for beginners than a bald list of Ancient Woodland Indicators (“A WIs’). Although this edition of ‘Rose’ is neither entirely suitable for the beginner nor for the advanced student it will be warmly welcomed by both amateur and professional botanists alike. Its combination of dichotomous keys and illustrations of diagnostic features bridges the gap between more elementary pictorial guides and more serious taxonomic works which can be so off-putting to botanists with little or no formal taxonomic training. This new edition is more comprehensive, stylish, and user-friendly than the original and is therefore likely to be the most popular field guide on the market for many years to come. K. J. WALKER REFERENCES COLEMAN, C. (2006). Wild Flower Key 2nd Edition: corrigenda and addenda, March 2006. BSBI News 102:71. PRIMAVESI, A. (2006). Don’t blame me. I never said it. BSBI News 103: 49-50. ROSE, F. (1989). Colour identification guide to grasses, sedges, rushes and ferns of the British Isles and north-western Europe. Viking, London. STACE, C. A. (1997). New flora of the British Isles 2nd ed. Cambridge University Press, Cambridge. STACE, C. A. (1999). Field flora of the British Isles. Cambridge University Press, Cambridge. Watsonia 26: 417-418 (2007) 417 Obituary LARCH SYLVIA GARRAD (1936-2005) With the death of Larch Garrad on 7 July 2005 the Society has lost its long-serving Recorder for Vice-county 71 and the Isle of Man, an authority of exceptional breadth on both its natural and local history. An only child, born in 1936 Larch’s closeness to her civil servant father ensured that from an early age she acquired both his voraciousness for reading and a keen interest in the details of the countryside, to the latter of which her enrolment in the Wild Flower Society gave a botanical slant. Brought up in Somerset, she attended the Grammar School for Girls in Weston-super-Mare before going on to Birmingham University to read Archae- ology and Ancient History. One of her archaeology lecturers happened to hail from the Isle of Man and it was through working on a student excavation there under his direction that she came to be acquainted with the Island. Meanwhile, excavation had taken her also to Greece, where the study of pottery styles appealed to her taxonomic bent and in due course became the subject of postgraduate research that culminated in the award of a doctorate. However, the career as an archae- ologist that ought to have followed turned out to be elusive: posts in the subject were then still very few and fiercely competed for, and eventually she had to settle for one as a librarian in Gloucestershire, the great distance of which from the Isle of Man involved fear- somely arduous and complicated journeys whenever she returned to see her parents, who were now living there in retirement. Providentially, a vacancy arose not long afterwards on the staff of the Manx Museum in Douglas — though for someone to take special responsibility for natural history, not in archaeology (which was well covered already). However, small museums necessarily prize broad interests and versatility above deep knowledge in one special field and she was able to gain the job on the basis of her reasonable familiarity with the Island’s natural history and its flora in particular. By that time, the compiling of a much- needed book-length account of the Island’s vascular plants had been under way for thirteen years, but one by one the few resident contributors of records had been dropping away through death, incapacity or removal ‘across’, and intensive fieldwork had largely fallen to occasional irruptions by visiting botanists that were all too brief. The arrival of a resident one at last who brought to this project personal enthusiasm as well as professional commitment was thus providential. Within two years, what is more, Larch struck up a friendship at evening class with a local schoolteacher nearing retire- ment who fancied using her impending greater leisure to pursue a long-existing interest in plants with greater seriousness. Marjorie Devereau supplied not only an extra pair of eyes, but two exceptionally sharp ones, which were to produce a string of increasingly startling discoveries in the years that followed; her better sight also enabled her to master sections of the flora, most notably Carex, that were unavoidably barred to Larch. As a car- Owner, moreover, she rescued the bus- dependent Larch from over-localised explor- ation. The innumerable additional records that arose from their joint efforts put paid to what had threatened to be indefinite deferment of the intended published volume, and the completion of that in 1984 owed an enormous amount to their cajoling and, ultimately, very full invo- lvement in the editorial process. As is often the way, the publication of the Flora had the paradoxical effect of lowering activity instead of sustaining or even increasing it. For lack of a concrete objective of comp- arable scope in its place the previous torrent of records fell off to a trickle, and competing interests filled the resulting vacuum. In Larch’s case there had long been no shortage of those and to collecting and recording non-marine molluscs and woodlice for the respective national atlases of those groups she has soon added investigating the uses of local plants as folk cures, establishing the identity of subfossil bones yielded by past excavations, co- © authoring a book on the Island’s industrial archaeology, and another on British holiday souvenirs. Conservation concern having mean- while spread to the Island very belatedly, the establishment of nature trails and promoting the work of a local wildlife trust made additional calls on her energies. 418 OBITUARY These multifarious involvements and the wide perspective they imparted made Larch the obvious person to contribute the relevant volume in the ‘Regional Naturalist’ series instituted by that transient shooting star of the British publishing scene, David & Charles. Once The Naturalist in the Isle of Man was off her hands, in 1972, they were busy again with an intended similar book on the Canary Islands, after a captivating holiday soaking herself in the natural history of those in_ turn. Unfortunately, though, that one had to be abandoned when the publisher ran _ into financial difficulties and had to cancel the contract. At the same time it was a symptom of widening horizons: concentrating so exclusively on one comparatively small island had begun to feel too restricting. Attendance at B.S.B.1. field meetings elsewhere, including on the Continent, had given her an appetite for other and richer floras. The limited prospects for professional advancement in so small and static a community, moreover, were meanwhile leading her to give thought to strengthening her qualifications and thus her geographical options. To that end, halfway through her career, she registered on an Open University course in botany. An early by-product of that was an impressive biometric analysis of a relic oak population on the Island’s cliffs. In later years the onset of heart trouble made solitary fieldwork medically inadvisable. Friends who relished the wonderful volleys of sometimes hilariously ill-assorted items of information for which she was well-known — and which found equal expression in her letters — were more than happy to have such an outstandingly well-informed guide on drives around the Island, but Larch was greatly restricted thenceforward in what she could accomplish. In her capacity as B.S.B.L Recorder she had diligently put together the relevant local data for the Society's Rare Species Survey, but she regretfully had to recognise that it had become physically beyond her to fulfil the recording requirements of the New Atlas. What she was able to supply on the other hand, was an exceptionally full set of records for the alien species, as she had for some years been concentrating on those with a view to publishing a supplement to the 1984 Flora. The unusually numerous exotics that flourish in the Island’s mild climate, trad- itionally disdained by recorders of the wild flora, had particularly engaged the attention of Marjorie Devereau and herself on their each acquiring gardens of their own and turning into enthusiastic plantswomen. From marrying archaeology came in 1985 A History of Manx Gardens and Gardening..., a publication on that then surprisingly almost virgin subject that — was so well-received than an enlarged edition was called for in 2003. The discovery in the meantime near Peel of a remarkable assembl- age of unusual adventives fired her to study those intensively as well, the preliminary results appearing in the Island’s principal natural history journal two years later, At the time of her death she was preparing for publication a full-scale Alien Flora of the Island, and hopefully that will have reached a sufficiently advanced state for that intention to be realised. Larch’s irrepressible vivacity, the product of a quick and very sharp mind, will be widely missed. Thankfully, a great part of the very extensive and remarkable diverse body of knowledge that she built up over rather more than forty ceaselessly active and copiously productive years, has been committed to print. The invaluable co-ordinating role that she performed in the field of Manx Studies will nevertheless be very difficult to replace. DAVID ALLEN SAND = Salles a aed etc IE — he Seat a NS t i) So a BRWN rR N 000 MOON NYY YYNYNN ABRWONEOCOOAINIDAARWHY Jelile H2. H3. H4. HS! H6. Lele H8. H9. H10. Je lltiie H12. ES: H14. ee Naess) en (Cag Ss 2) West Cornwall Scilly East Cornwall South Devon North Devon South Somerset North Somerset North Wiltshire South Wiltshire Dorset Isle of Wight . South Hampshire North Hampshire West Sussex East Sussex East Kent West Kent Surrey ‘South Essex North Essex Hertfordshire . Monmouthshire . Glamorgan . Brecknockshire . Radnorshire . Carmarthenshire . Dumfriesshire . Kircudbrightshire Wigtownshire . Ayrshire . Renfrewshire . Lanarkshire . Peeblesshire Selkirkshire . Roxburghshire . Berwickshire . East Lothian . Midlothian West Lothian . Fife &Kinross South Kerry North Kerry West Cork Mid Cork East Cork Co. Waterford South Tipperary Co. Limerick Co" Clare North Tipperary Co. Kilkenny Co. Wexford Co. Carlow Laois NAMES OF VICE-COUNTIES IN WATSONIA 7M Jpeg Ppy. 24. DS) 26. Jai 28. ray); 30. Sil, 3s. 33: 34. 36. Sift 38. Sy), 40. 3)3h 45. 46. 47. 48. 49. 86. 87. 88. 89. 90. 91. 92. 93) 94. 25): 96. 96b. OT. 98. H15. H16. laBLVe H18. jl). H20. HZ H22. H23) H24. H25. H26. 27: H28. S (113) Channel Islands ENGLAND Middlesex 54. Berkshire 5)5)- Oxfordshire DDD: Buckinghamshire 56. East Suffolk ay West Suffolk 58. East Norfolk 59. West Norfolk 60. Cambridgeshire 61. Bedfordshire 62. Huntingdonshire 63. Northamptonshire 64. East Gloucestershire 65. West Gloucestershire 66. Herefordshire 67. Worcestershire 68. Warwickshire 69. Staffordshire 69b. Shropshire 70. South Lincolnshire JAY WALES Pembrokeshire 50. Cardiganshire ay) ls Montgomeryshire 5, Merioneth Caernarfonshire SCOTLAND Stirlingshire 99. West Perthshire 100. Mid Perthshire sm OL East Perthshire 102. Angus 103: Kincardineshire 104. South Aberdeenshire 105. North Aberdeenshire 106. Banffshire 107. Moray 108. Easterness 109. Nairnshire 110. Westerness MUL Main Argyll Ps. IRELAND South-east Galway F129} West Galway H30. North-east Galway Heike Offaly H32. Co. Kildare 33) Co. Wicklow H34. Co. Dublin H35. Meath H36. Westmeath H37. Co. Longford H38. Co. Roscommon H39. East Mayo H40. West Mayo Co. Sligo North Lincolnshire Leicestershire Rutland Nottinghamshire Derbyshire Cheshire South Lancashire West Lancashire South-east Yorkshire North-east Yorkshire South-west Yorkshire Mid-west Yorkshire North-west Yorkshire County Durham South Northumberland North Northumberland Westmorland Furness Cumberland Isle of Man Denbighshire Flintshire Anglesey Dunbarton Clyde Isles Kintyre South Ebudes Mid Ebudes North Ebudes Wester Ross Easter Ross East Sutherland West Sutherland Caithness Outer Hebrides Orkney Shetland Co. Leitrim Co. Cavan Co. Louth Co. Monaghan Fermanagh East Donegal West Donegal Tyrone Co. Armagh Co. Down Co. Antrim Co. Londonderry INSTRUCTIONS TO CONTRIBUTORS Scope: Authors are invited to submit Papers and Notes concerning British and Irish vascular plants, their taxonomy, biosystematics, ecology, distribution and conservation, as well as topics of a more general or historical nature. Authors should consult the Hon. Receiving Editor for advice on suitability or any other matter relating to submission of manuscripts. Papers and Notes must be submitted in duplicate, typewritten on one side of the paper, with wide margins and double-spaced throughout. Pages should be numbered. Submission of final edited copy on computer disc will be requested, but two hard copies of the text are acceptable if computer . facilities are not available. Format should follow that used in recent issues of Watsonia. Underline where italics are required. Names of periodicals should be given in full, and herbaria abbreviated as in British and Irish herbaria (Kent & Allen 1984). The Latin names and English names of plants should follow the New Flora of the British Isles (Stace 1997). Further details on format can be obtained from the Hon. Receiving Editor or by viewing the website at: http://www.bsbi.org.uk/new_style_manual.htm Tables, figure legends & appendices should be typed on separate sheets and attached at the end of the typescript. Figures should be drawn in black ink or be laser-printed and identified in pencil on the back with their number and the author’s name. They should be no more than three times final size, bearing in mind they will normally be reduced to occupy the full width of a page. Scale-bars are essential on plant illustrations and maps. Lettering should be of high quality and may be done in pencil and left to the printer. Black and white photographs can be accepted if they assist in the understanding of the article. If you are able to submit figures on disc please contact the Receiving Editor to check they are in a suitable format. Contributors must sign a copyright declaration prior to publication which assigns the copyright of their material to the Botanical Society of the British Isles. Twenty-five offprints are given free to authors of Papers and Notes; further copies may be purchased in multiples of 25 at the current price. The Society takes no responsibility for the views expressed by authors of Papers, Notes, Book Reviews or Obituaries. Submission of manuscripts Papers and Notes: Mr M. N. Sanford, c/o The Museum, High Street, Ipswich, Suffolk, IP] 3QH. Books for Review: Dr C. D. Preston, Biological Records Centre, Monks Wood, Abbots Ripton, Huntingdon, PEI7 2LS. Plant Records: the appropriate vice-county recorder, who should then send them to Mike Porter, 5 West Avenue, Wigton, Cumbria CA7 9LG. Obituaries: Mrs M. Briggs, 9 Arun Prospect, Pulborough, West Sussex, RH20 IAL. Back issues of Watsonia are available from the official agents for BSBI Publications, Summerfield Books (John & Sue Atkins), Summerfield House, High Street, Brough, Kirkby Stephen, Cumbria CA17 4AX. Tel: 017683 41577. Fax: 017683 41687. E-mail: bsbipubs @ beeb.net Watsonia February 2007 Volume twenty six Part three S. M. Walters Memorial Issue Contents SANFORD, M. N. Introduction =. OBITUARY AND BIBLIOGRAPHY Stuart Max Walters ( 1920-2005) a KADEREIT, J. W. & WESTBERG, E. Determinants of phylogeographic structure: a comparative study of seven coastal gee. oe saan across their European range ' : GURNEY, M., PRESTON, C. D., BARRETT, J. & Brigas, D. ecnamien bee Oxlip Primula elatior (L.) Hill and Primrose P. vulgaris Hudson, and the identification of their variable hybrid P. xdigenea A. Kerner PRESTON, C. D. Which vascular plants are found at the northern or Cane edge of their European range in the British Isles? PEARMAN, D. A. ‘Far from any house’ — assessing the status of doubly native species in the flora of the British Isles WALKER, K. J. The last sae five years: recent changes in the . flo of ie British Isles .. BEECROFT, R. C., ene, C. i & MOUNTFORD, J. o Water Gennes Teucrium scordium L. in Cambridgeshire: back from the brink of extinction ... SWAN, G. A. & RICHARDS, A. J. The Eleocharis mamillata H. Lindb. fil. pec (Cyperaceae) in the British Isles :; MIDDLETON, R. & MIDDLETON, J. R. The Tee of Hull Bese ‘ PAGE, C. N., MCHAFFIE, H. & BUTLER, J. K. A new far northern hybrid iene from Scotland: Equisetum fae id 2, CaN: me au ey JE, x E. pratense Ehrh.) see : GRANADOS, L. & LANE, S. D. A fine ite bade of ici enuanonnnenen ea floristic parameters in three populations of eae vigursil pee a rare annual endemic to Devon and Cornwall PASHLEY, C. H., BAILEY, J. P. & FERRIS, C. Clonal joa in 1 British sopulaeene of the alien invasive Giant Knotweed, Fallopia sachalinensis (F. Schmidt) Ronse Decraene, in the context of European and Japanese plants ae EL-BAKATOUSHI, R., RICHARDS, A. J. & WOLFF, K. Introgression between Plantago major L. subspecies ee, and eee intermedia ae mae in a British population TENNANT, D. J. Names and een oes ee ses sallow “(Salis L) yb discovered recently in Scotland Teh Posse | ee sae NOTES Foley, M. J. Y. The Gunna (v.c. ee record for Carex tiie Schum. (Cyperaceae) Pearman, D. A. & Biinaide. B. Vaterianeney eriocarpa Desv. as a aiscniiede native in Britain — an update... Allen, D. E. Rubus milesianus wholly a garden escape in Britain? PLANT RECORDS BOOK REVIEWS OBITUARY ISSN 0043-1532 Typeset by D. K. & M. N. SANFORD Printed in Great Britain by PALLADIAN PRESS, UNIT E, CHANDLERS ROW, PORT LANE, COLCHESTER, ESSEX CO1 2HG Winn nii 213 215-227 229-238 239-251 253-269 271-290 291-302 303-316 317-325 327-338 339-345 347-358 359-371 373-379 381-383 385-386 387 388-389 391-407 409-415 417-418 ON wad ATeUNIA B07 Journal of the Botanical Society of the British Isles Volume 26 Part 4 August 2007 Editors: M. Briggs, M. J. Y. Foley, D. L. Kelly, D. R. McKean, M.S. Porter, M. N. Sanford Botanical Society of the British Isles Registered Charity Number: 212560 Applications for membership should be addressed to the Hon. General Secretary, c/o Department of Botany, The Natural History Museum, Cromwell Road, London, SW7 5BD, from whom copies of the Society’s Prospectus may be obtained. Officers for 2007—2008 President, Dr R. J. Gornall Vice-Presidents, Prof. R. M. Bateman, Miss L. Farrell, Mr R. G. Woods, Dr F. J. Rumsey Honorary General Secretary, Mr D. A. Pearman Honorary Treasurer, Mr M. E. Braithwaite Editors of Watsonia Papers and Notes, M. J. Y. Foley, D. L. Kelly, D. R. McKean, M. N. Sanford* Book Reviews, J. Atkins Plant Records, M. S. Porter Obituaries, M. Briggs *Receiving editor, to whom all MSS should be sent (see inside back cover). © 2007 Botanical Society of the British Isles The Society takes no responsibility for the views expressed by authors of Papers, Notes, Book Reviews or Obituaries. The cover illustration of Euphorbia hyberna L. (Irish Spurge) was drawn by Rosemary Wise. Watsonia 26: 419-432 (2007) 419 One-way introgressive hybridisation between Sorbus aria and S. torminalis (Rosaceae) in southern Britain Dei PRICE 43 Wonastow Road, Monmouth NP25 5DG and ies Ganicr: Department of Biodiversity and Systematic Biology, National Museum of Wales, Cardiff CF 10 3NP ABSTRACT Material of Sorbus aria-like plants from the Wye Valley with acutely-lobed leaves has _ been investigated using comparative morphology and reproductive fertility, which show that at least some such plants are S. aria introgressed with S. torminalis. The introgression was found to occur only from S. aria into S. torminalis. The primary and introgressed hybrids have lower pollen and fruit fertility than the parents. The parents show wide- spread overlap in distribution in southern Britain but introgression has been confirmed only for the Wye Valley. Sorbus =< vagensis should be treated as a hybrid, not as a species. KEYWORDS: Fertility, morphology, pollen, Sorbus latifolia, Sorbus x tomentella, Sorbus < vagensis. INTRODUCTION The Sorbus latifolia (Lam.) Pers., Broad-leaved Whitebeam aggregate comprises taxa which have arisen by hybridisation between S. torminalis (L.) Crantz, Wild Service-tree, and members of the S. aria (L.) Crantz, Common Whitebeam aggregate (Warburg 1962; Sell 1989; Nelson-Jones ef al. 2002). They are characterised by having leaves with triangular, acute or acuminate lobes and greenish-grey tomentum below, and large fruits which are brown, orange or yellow when ripe. In Britain, the S. latifolia aggregate was first separated by Syme (1875, as Pyrus latifolia Syme), and now contains seven taxa (Sell 1989), six of which are apomictic polyploids (S. bristoliensis Wilm., S. croceocarpa P. D. Sell, S. decipiens (Bechst.) Irmisch, S. devoniensis E. F. Warb., S. latifolia sensu stricto and S. subcuneata Wilm.), and the other is the primary diploid hybrid S. x vagensis Wilm. *e-mail: tim.rich@museumwales.ac.uk When he first described it, Wilmott (1934) considered that S. x vagensis might be S. tor- minalis x anglica Hedl. or x porrigentiformis E. F. Warb., and suggested it grew true from seed after noting the morphological uniformity of a cluster of trees a little south of Symonds Yat top. He chose the epithet to reflect the occurrence of the hybrid in the Wye Valley (of the river Wye or flumen Vaga). Warburg subsequently found that the uniformity noted by Wilmott was due to suckering rather than reproduction from seed (Riddelsdell ef al. 1948), and he treated it as a variable, diploid hybrid between S. aria and S. torminalis (Warburg 1952), which is now its accepted parentage. It is now known to have arisen a number of times independently, with S. torminalis as the female parent (Proctor & Groenhof 1992; Lemche 1999; Fay et al. 2002; Chester et al. 2007). Sell (1989) treated S. x vagensis as a species rather than a hybrid since British material did not match in morphology continental material of the same parentage; we follow his nomenclature here but not his concept of the taxon as a species. During a series of Sorbus surveys of cliffs in the Wye Valley in 2000-2003 commissioned by English Nature (now Natural England), L. Houston, C. Charles and A. C. Tillotson repeat- edly collected forms of S. aria with acutely- lobed leaves (Houston et a/. 2001, 2002, 2003, 2004; Figure la, b). In exasperation at the absence of any name to call them, they were nicknamed Sorbus “Yet another form” by C. Charles. These plants were shown to T. Rich, who, after much discussion, began to suspect these might be S. aria introgressed with S. torminalis, not previously reported from Britain but known from the continent (Chabert 1906; Aas et al. 1994; Oddou-Muratorio et al. 2001). 420 Dia PRICE Sail CaGe RICH TABLE 1. CHARACTERS DISTINGUISHING SORBUS ARIA, S. TORMINALIS AND THEIR PRIMARY DIPLOID HYBRID S. x VAGENSIS, COMPILED FROM THE LITERATURE (E. G. WARBURG 1962, CHALLICE & KOVANDA 1978) AND PERSONAL OBSERVATION Habit No suckering Weakly suckering Strongly suckering Petiole length 7—24(-29) mm (15—)20-35(-43) mm (15—)20—50(—S8) mm Leaf shape +Orbicular, ovate, elliptical Ovate, elliptical, oblong or Ovate, 0-8—1-3(—1-5) times as or oblong (rarely obovate), | rhombic-elliptical, (0.8—) long as wide (1-0—)1-1—2:0(-2:4) times as__1-0—1-5(—1-9) times as long long as wide as wide Leaf veins (6—)9—14(—15) pairs 6—10(—12) pairs (3—)4—6(—7) pairs Leaf pubescence White tomentose below Greenish-grey tomentose + Glabrous below at maturity below, sometimes subglabrous Leaf apex Obtuse to acute Acute or subacute Acuminate Leaf base Truncate to rounded (rarely Rounded or cuneate Rounded, truncate or cordate, cuneate or cordate) rarely broadly cuneate Leaf lobing Unlobed or with shallow Shallow lobed with acute or Deeply lobed with triangular, — broadly rounded lobes to 1/6 acuminate lobes to '%4 way — acute or acuminate lobes to 4 way to midrib to the midrib (rarely all) way to the midrib Leaf margin Doubly crenate-serrate, teeth Finely serrate with small Finely serrate with small with one side a little longer somewhat appressed teeth — strongly asymmetric than the other appressed teeth Inflorescence Densely crowded Crowded Lax Sepals Eglandular Sparsely glandular Glandular Styles 2-3, free, lanuginous at base 2-3, variably joined upto 3, joined up to 2—4Away, Fruit colour Lenticels on fruits Sorbus aria Red Few to medium, small to medium lenticels, sometimes Sorbus < vagensis Sorbus torminalis half way, lanuginous at base Yellow, orange or glabrous at base Brown brownish-orange Medium to numerous, small Numerous small lenticels and and medium lenticels large ones Fruit length (9—)10—15(—16) mm (9-)10-17 mm In this paper we present the results of morphological and reproductive biology investigations into these Wye Valley S. aria plants with acutely-lobed leaves. Sorbus aria, S. torminalis and S. x vagensis can be readily separated morphologically using many characters (Table 1), and putative backcrosses might be expected to show some characters intermediate between S. x vagensis and S. aria, such as relatively long petioles, few pairs of veins and greener undersides to the leaves. Hybrids are often sterile or show reduced fertility compared to their parents. In order for back-crossing to occur, some of the primary hybrids and backcrosses must produce at least some fertile pollen and/or seeds. The primary hybrids were reported as either sterile or fertile in France (de Poucques 1951), but this has not been studied in Britain so the relative fertility of the taxa has been investigated from pollen and fruit. some large ones (9-)10-17(-18) mm We have also compiled information on the distribution of S. < vagensis and the locations where its parents occur together to see how widespread back-crossing might potentially be in Britain. METHODS COMPARATIVE MORPHOLOGY Herbarium material of S. aria (21 samples), S. torminalis (18 samples), S. =< vagensis (19 samples) and putative backcrosses of S. x vagensis with S. aria (13 samples) were selected from NMW (Appendix 1). Material was selected from the Wye Valley and from elsewhere in Britain where hybrids are unknown. Some _ material previously investigated for DNA analysis (Fay ef al. 2002) was also selected (marked in Appendix 1). The following set of eight characters were selected SORBUS RESULTS It was possible to name most of the plants using existing Asperula accounts, but four British specimens were intermediate between subsp. cynanchica and subsp. occidentalis and could not be named with confidence. Material of subsp. occidentalis from Spain and France appeared to be larger and distinct from material called subsp. occidentalis in the British Isles. With few exceptions, scoring of most of the characters was straightforward. It was often difficult to decide if the lateral ‘stems’ without leaves were stolons or rhizomes; where these organs were obviously underground in the original situation, they lacked adventitious roots and were probably best regarded as subterranean stolons rather than rhizomes. The stolons were most prevalent in mat-forming plants ascribed to subsp. occidentalis, but the colour was also quite variable and inconsistent, and whilst some roots of subsp. occidentalis were strongly and distinctively orange-coloured, some were not. Some underground stolons also VARIATION IN ASPERULA CYNANCHICA 435 Component 2 Component | FIGURE 1. PCA of Asperula cynanchica taxa. X, A. cynanchica, British Isles and Europe. @, Subsp. occidentalis, Spain/France. O, ‘Subsp. occidentalis’, British Isles. H, A. cynanchica var. densiflora sensu Sell & Murrell, British Isles. +, Intermediate between 4. cynanchica and A. occidentalis, British Isles. clearly bore leaves at some stage, and had presumably been buried by sand or soil, resulting in further upward growth. We do not regard the stolons as providing reliable taxo- nomic characters. The corolla appendage was not always well defined, even when corollas were soaked in hot water and laid out for careful examination. The Principal Components Analysis of the all taxa is shown in Figure 1. Axis 1 accounts for 28% of the variation and is the most important for separating the taxa, the major vari- ables being (in decreasing order of importance): middle leaf length, upper leaf length:width ratio, middle leaf length:width ratio, upper leaf length and length of middle internode. Axis 2 accounts for 13% of the variation but provides little separation between the taxa, the main variables being the lower leaf length:width ratio, and the upper leaf length to widest point. Axis 3 accounts for 9% of the variation and contributed little to separating the taxa. There is a strong pattern to the variation in Figure 1. Material of Spanish and French subsp. occidentalis, including material from the type locality at Biarritz, forms a reasonable cluster 436 Re BIC ODD Sale CGA RICE in the top centre, largely distinct from British and Irish material named subsp. occidentalis on the left, but overlapping to a degree with subsp. cynanchica (British, Irish and _ continental Europe) on the right. The outlier to the left is a small, dense plant from Biarritz, but is otherwise similar to other subsp. occidentalis from the same locality. There is good separation of British and Irish material named subsp. occidentalis from subsp. cynanchica. There is complete overlap of British and Irish subsp. cynanchica var. densiflora with British and Irish material named subsp. occidentalis. The intermediate plants from England are largely in an area of overlap at the bottom of the figure between subsp. cynanchica and British and Irish subsp. cynanchica var. densiflora and subsp. occidentalis. Separate Principal Components Analyses comparing subsp. cynanchica with British and Irish subsp. occidentalis, and comparing French/Spanish subsp. occidentalis with British and Irish subsp. occidentalis show better separation compared to Figure | (as might be expected for simpler data sets), whilst French/ Spanish subsp. occidentalis still cluster within the variation in subsp. cynanchica when compared separately (data not presented). All plants flowered and fruited freely in culti- vation. The subsp. occidentalis plants retained their prostrate, dense habit and looked very similar to the wild plants, other than becoming slightly larger. Both subsp. cynanchica plants became larger and sprawling. DISCUSSION The morphological analyses indicate that there are three taxa which differ mainly in quantitative characters: subsp. cynanchica, subsp. occidentalis from France/Spain, and a third taxon which includes subsp. cynanchica var. densiflora and British and Irish material which has been called subsp. occidentalis. We do not accept that subsp. occidentalis occurs in the British Isles. Asperula cynanchica is a variable taxon, and it remains to be seen whether true subsp. occidentalis from France and Spain is best retained within 4. cynanchica or restored to a species in its own right. Its relegation to subspecific status was based on observations of material from both the continent and the British Isles (pers. comm. C. A. Stace 2007), but the latter part of that material we now refer to subsp. cynanchica var. densiflora. Although our Principal Components Analysis shows that subsp. occidentalis is nested within the variable subsp. cynanchica, it is based on relatively few specimens of subsp. occidentalis and the inclu- sion of further samples might improve its definition. Rouy (1903) described two varieties of A. occidentalis at Biarritz, a. galiiformis and B cynanchiciformis, the former small and dense, the latter more lax. This variation is also evident in the duplicates collected there later by E. J. Neyraut (LY, MA, P). These seem to be forms of the same taxon related to growth conditions, which could be verified by culti- vation experiments. We suggest that no further taxonomic changes are made until a wider review of A. cynanchica and its close relatives throughout Europe is undertaken. We also suggest that the name var. densiflora (as used by Sell & Murrell 2006) can be used for the third taxon (i.e. including British and Irish material which has been called subsp. occidentalis) until the full infraspecific variation in A. cynanchica has been reviewed. Var. densiflora was described by Grenier & Godron (1852) as having “Fleurs plus nom- breuses; tiges plus courtes, plus ramassées et plus étalées” [Flowers more numerous; stems shorter, more branched and prostrate], and they recorded its habitat as dunes of the Atlantic and Mediterranean coasts. This description fits the British and Irish plants reasonably well, but we have not attempted to trace types. Rouy (1903) also listed nine other varieties in France. Although var. densiflora maintains _ its growth form in cultivation, some populations look slightly different one to another, with Irish material in particular being somewhat variable. The South Wales material is relatively uniform. Our current feeling is that the dense, prostrate growth form may have originated a number of times from local A. cynanchica in response to the strong environmental selection pressure in coastal habitats, and is thus polyphyletic. This could be easily tested using molecular methods. Diagnostic descriptions of the taxa from our data are given below. Measurements are presented as (minimum—) 10 percentile—90 percentile (-maximum). VARIATION IN ASPERULA CYNANCHICA 437 Pie ae y) ae C ; FIGURE 2. Asperula cynanchica var. cynanchica silhouettes. A, Petersfield (v.c. 12). B, Babraham (v.c. 29). C, White Horse Hill (v.c. 22). Scale bar 1 cm. Asperula cynanchica L. subsp. cynanchica var. cynanchica (Fig. 2). Plant slender, lax, stems weakly ascending, stolons sometimes present, brownish. Lower stem leaves (2:5—)3—7(—13) mm x (0-3—)0-5— 1-2(-2:1) mm, (1-7—)3—13(-20) times as long as wide. Middle stem leaves (5-6—)7—19(—26) mm xX (0-3—)0-5—1-1(—2) mm, (4:6—)9—28(-45) times as long as wide. Middle internode (4—-) 8-6-27(-44) mm, (0-5—)0-9-2:4(-4-4) times as long as adjacent leaf. Upper stem leaves (S5—)8— 19-35) mm x (0-4-)0-5—1(-1-5), (6:-4—)10-32 (-50) times as long as wide. Inflorescence lax. Terminal pedicel (0—)0-4—1-1(—1-6) mm. Corolla tube (0-4—-)0-6—1-1(—-1-:5) mm, (0-3-) 0-4—0-7(—0-9) times as long as lobe. Corolla lobe (1—)1-3—2-2(-2°8) mm x (0-4—)0-5—1-1( 1-4), lobe (1:3—)1-6—3-2(-4:5) longer than wide. Lobe appendages usually present. Distribution: Widespread in Europe on calcareous grasslands and occasionally on dunes. The map in Preston ef al. (2002) shows its distribution in southern Britain and Ireland, though it is rarer in South Wales than var. densiflora. 438 R. B. CODD & T. C. G. RICH A © FIGURE 3. Asperula cynanchica vat. densiflora silhouettes. A, B. Whorlebury Hill (v.c. 6). C, Bembridge (v.c. 10). Scale bar 1 cm. FIGURE 4. Distribution of Asperula cynanchica var. densiflora in the British Isles. Asperula cynanchica L. subsp. cynanchica var. densiflora Gren. & Godr. (Fig. 3). Plant small, compact, often mat-forming, prostrate, stolons brown or brownish-orange, or absent. Lower stem leaves (1—)2:3-4:2(-4:6) mm xX (0:4—)0-6—1-4(—2:0) mm, (1-9—)2-1—5-7( 9-8) times as long as wide. Middle stem leaves (3:2—)4:2—-9(-14) mm x (0-4-)0-6—-1-4(-2) mm, (3—)4-11(-18) times as long as wide. Middle internode 3—12(—16) mm, (0-5—)0-6—1-5(-2°3) times as long as adjacent leaf. Upper stem leaves 4—10(-15) mm x (0-4—-)0-5—1-3(-1-7) mm, (4—) 5:-5-13(-19) times as along as_ wide. Inflorescence crowded. Terminal pedicel (0-2-) 0-3—-1-6(-1:7) mm. Corolla tube (0-8—)1—1-5(— 1-6) mm, (0-4-)0-6—1 times as long as lobe. Corolla lobe (1—)1-3—1-9(-2:3) mm x (0:4—)0-6— 1-3 mm, (1-2—)1-3—2-5(-2°8) times as long as wide. Lobe appendages distinct, weak or absent. VARIATION IN ASPERULA CYNANCHICA 439 FIGURE 5. Asperula cynanchica subsp. occidentalis silhouettes. A, Liencres, Spain. B, C, Biarritz, France. D, El Tyo, Spain. Scale bar | cm. Distribution: Southern Britain and Ireland (Figure 4), France, possibly elsewhere in Europe too. It 1s most frequent near the coast on sea cliffs and sand dunes, but may also occur inland. Some herbarium specimens contain a mixture of material suggesting that both varieties can grow together. Details of the specimens seen are given in Appendix 1. Asperula cynanchica L. subsp. occidentalis (Rouy) Stace (Fig. 5). Plant large, lax, stolons usually present, sometimes absent, bright orange or brown. Lower stem leaves (2:3—)2-8—5-1(—7-8) mm x 0-9-1-9(-2°6), (1:5—)2—5-5(—6:5) times as long as wide. Middle stem leaves (5-8—)8—11(—13°5) mm xX (0-7—)0-8—1-5(—1-8) mm, (3-9—)4-5—13(— 17) times as long as wide. Middle internode 5— 14(-17) mm, 0-6—1-4 times as long as adjacent leaf. Upper stem leaves (7:6—)9-15-3 mm x 0-7—1-1(-1-5), (7—)9-17(-21) times as long as wide. Inflorescence lax. Terminal pedicel 0-0-3 (—0-4) mm, often very short. Corolla tube 0-9— 1-2(-1:3) mm, 0-5—0-8 times as long as lobe. Corolla lobe (1:2—)1-4—2:1 mm x 0-4—-1-1 mm, 1:5-3-6 times as long as wide. Lobe appendages always present. 440 Re Ba CODD sails CxGaRiC@H Distribution: France (Basses-Pyrenees only; Ruderon (1995)) and northern Spain (Asturias, Cantabria, Guipuzcoa, Vizcaya) on calcareous grasslands and dunes. The specimens seen are cited in Appendix 2. We have not seen voucher material of the records on the French flora website (www.tela- botanica.org) for Normandy but suspect they are likely to refer to var. densiflora, and the specimen from Le Poulignes, Loire-Inferior (P) is var. densiflora. We have no reason to doubt records cited in the Spanish flora website (Poryecto Anthos: www.anthos.es) for northern Spain as they have been revised recently for Flora Iberica. ACKNOWLEDGMENTS We are grateful to the Keepers of CGE, DBN, MA, P and RNG for access to or loan of material, and to Gaétan Guignard (LY) for images of the Rouy types, and Jerome Sawtschuk for help with translation. The map was plotted using DMAPW by Alan Morton. KEY il. Middle stem leaves (5—)7—19(—26) mm long, (5—)9—28(-45) times as long as wide and (0:5—)0:9-2:4(-4-4) times as long as adjacent leaf .......... subsp. cynanchica var. cynanchica Ile Middle stem leaves (3—)4—11(—14) mm long, 4—-13(—18) times as long as wide and (0-5—) 0-6—2°3) times as long as adjacent leaf .....-.g2.0...scccscecerescocosceuee cece eeeee ee eee 2, De Terminal pedicel (0-2—)0-3—1-6(—1-7) mm; corolla lobe appendages distinct, weak or absent Fe Pr ce nt Re ee EMP Cc Son Mie be Goh sofidasionds subsp. cynanchica var. densiflora De Terminal pedicel 0—0-3(—0-4) mm, often very short; corolla lobe appendages distinct ......... sian qadaseGeMeshuas a deen qnsonneaaidsincasmgan teats odes ue te ee see ae mene Seance Shc ete eae ee See subsp. occidentalis REFERENCES CLAPHAM, A. R., TUTIN, T. G. & Moore, D. M. (1987). Flora of the British Isles. 3rd edition. Cambridge University Press, Cambridge. CONTEJEAN, M. Ch. (1865). Note sur quelques plantes rare ou critique du midi de la France. Bulletin de la Société Botanique de France 12: 217-221. EHRENDORFER, F. & KRENDL, F. (1976). Asperula L., in TUTIN, T. G., HEYWOOD, V. H., BURGES, N. A., Moore, D. M., VALENTINE, D. H., WALTERS, S. M. & WEBB, DD. A. (1976). Flora Europaea 4: 4-14. Cambridge University Bressy © ambridge. GRENIER, J. C. M. & GODRON, D. A. (1852). Flore de France, 2. E. Savy, Paris. HAMMER, @., HARPER, D. A. T. & RYAN, P. D. (2001). PAST: Paleontological Statistics Software Package for Education and Data Analysis. Palaeontologia Electronica 4(1): 9pp. http://palaeo- electronica.org/2001_ 1/past/issuel_01.htm. PRESTON, C. D., PEARMAN, D. A. & DINES, T. D. (2002). New Atlas of the British and Irish flora. Oxford University Press, Oxford. Ricu, T. C. G. & JERMY, A. C. (1998). Plant Crib 1998. Botanical Society of the British Isles, London. Rouy, G. (1903). Flore de France. Volume 8. Asnicres, Paris & Rochefort. RUDERON, J.-P. (1995). Inventaire des plants protégées en France. Editions Nathan, Paris. SELL, P. D. & MURRELL, G. (2006). Flora of Great Britain and Ireland. Volume 4. Campanulaceae to Asteraceae. Cambridge University Press, Cambridge. STACE, C. A. (1989). New combinations in the British and Irish flora. Watsonia 17: 444. STACE, C. A. (1997). New flora of the British Isles, 2" ed. Cambridge University Press, Cambridge. TUTIN, T. G. & CHATER, A. O. (1974). Asperula occidentalis Rouy in the British Isles. Watsonia 10: 170-171. WEBB, D. A. & SCANNELL, M. J. P. (1983). Flora of Connemara and the Burren. Royal Dublin Society and Cambridge University Press, Cambridge. (Accepted May 2007) VARIATION IN ASPERULA CYNANCHICA 44] APPENDIX 1 BRITISH AND IRISH SPECIMENS OF ASPERULA CYNANCHICA VAR. DENSIFLORA SEEN ENGLAND Worlebury Hill (V.c. 6), 14 Aug 1887, J. W. White (CGE, NMW). Worle Hill, Weston Super Mare (v.c. 6), June 1842, G. W. H. Thwaites (CGE). Marlborough (v.c. 7), 8 July 1885, E. S. Marshall, (CGE). Kingsdown [Kingston?] (v.c. 9), 15 June 1899, E. F. Linton (NMW). Portland (v.c. 9), 1837, A. Bloxham (CGE). Church Ope Cove, Portland (v.c. 9), 6 July 2000 and 11 August 2003, T Rich (NMW). Ventnor (v.c. 10), August 1834, F. Ballard (NMW). Golf links at sea level, Bembridge (v.c. 10), 12 July 1905, Miss C. Bickham (CGE). Bembridge (v.c. 10), July 1829, W. Annesley (CGE, left hand specimen). Short turf near cliff edge, Beachy Head (v.c. 14), July 1955, F. Rose (NMW). WALES Near Swansea (v.c. 41), 1835, T. A. Babington (CGE). Nottage (v.c. 41), 1891, A. H. J. (NMW). Near Rhossili (v.c. 41), 25 May 1953, B. A. Miles (CGE). Rhossili cliff (v.c. 41), 28 May, 1929 (NMW). On limestone on Gower (v.c 41), June 1835, T. A. Babington (CGE). Langland Cliff, Gower (v.c. 41), 16 June 1928, V. M. Peel (NMW). Mumbles Hill (v.c. 41), 6 June 1993, G. Hutchinson (NMW). Mumbles Head (v.c. 41), 4 June 1997, T. Rich (NMW). Horton Dunes, Gower (v.c. 41), 28 June 1945, J. A. Webb (NMW), and undated, Anon (NMW). Limestone cliffs between Langland and Caswell Bay (v.c. 41), 30 May 1909, G. R. Willan (NMW). Whitford Burrows (v.c. 41), July 1993, P. Russell (NMW). Porthcawl (v.c. 41), 1905, A. H. J. (NMW), 7 June 1901, W. A. Shoolbred (NMW) and 17 June 1931, A. E. Wade (NMW). Kenfig, 3 July 1982, H. J. M. Bowen (RNG). Pennard Castle (v.c. 45), June 1920, E. N. Thomas (NMW), and 1971, R. A. Boniface (NMW). Pembrey Local Nature Reserve (v.c. 44), 5 July 2000, R. D. Pryce & K. A. Cottingham (NMW). Carreg Cennan (v.c. 44), 15 July 1945, J. A. Webb (NMW). Pembrey (v.c. 44), 8 July 1936, J. M. Lewis (NMW). Laugharne Burrows (v.c. 44), 1987, J. Rees (NMW). Dry dunes, Laugharne Burrows (v.c. 44), 4 July 1982, R. D. Pryce (NMW). Sand dunes, Burry Port (v.c. 44), 13 July 1967, J. R. Gates (NMW). Pendine Dunes (v.c. 44), 28 June 1952 (NMW). Whitesand Bay (v.c. 45), 5 April 1975, R. M. Burton (NMW). Cliff, Barafundle Bay (v.c. 45), 25 July 1960, A. C. Powell (NMW). Wall on the Burrows, Tenby (v.c. 45), 1881, J. Sidebotham (CGE). Tenby (v.c. 45), 19 August 1848, C. C. Babington (CGE). Tenby Burrows, 2 July 1931, E. C. Howells (NMW). Giltar Point, 12 July 1999, T. Rich (NMW). Lydstep, 12 July 1999, T. Rich (NMW). Saddle Point (v.c. 45), 23 July 1999, T. Rich (NMW). Angle (v.c. 45), 1871, illeg. (CGE). Sand dunes, Penally, 2 August 1975, J. E. Lousley (RNG). IRELAND Smerwick Cove (v.c. H2), 13 July 1841, C. C. Babington (CGE). Abundant on the sandhills, Castlegregory (v.c. H2), 24 June 1902, E. S. Marshall (CGE, NMFW). Sandhills, Lehinch (v.c. H9), 8 August 1893, H. C. Levinge (DBN) and July 1880, R. M. Barrington (DBN). Muckinish, Ballyvaughan (V.c. H9), 23 August 1895, N. Colgan (DBN, sand hill form). Near Kilmnovie, Aran Isles (v.c. H9), 5 September 1976, M. Scannell (DBN). Ballybunion (v.c. H16), August 1866, M. Barrington (DBN). Bunowen penisular (H16), 16 June 1974, M. Scannell (DBN). Sandy ground, Mannim (v.c. H16), September 1979, M. Scannell (DBN), and 23 August 1974, D. McClintock (RNG). Dogs Bay (v.c. H16), 1966, F. Rose (NMW). Roundstone (v.c. H16), undated, R. M Barrington (DBN). Near the beaches near Roundstone (v.c. H16), 12 September 1873, C. C. Babington (CGE). 442 Re BI CODDica I C1GsRIGH APPENDIX 2 SPECIMENS OF ASPERULA CYNANCHICA SUBSP. OCCIDENTALIS SEEN FRANCE Biarritz, cote des basques, entre parents, 18 September 1869, M. Ch. Contejean (LY, holotype of var. galiiformis); sables maritime, May 1878, Bordere, (LY, holotype of var. cynanchiciformis); coastal sand dunes, 9 August 1903, E. J. Neyraut (LY, MA nos. 117219 and 117218, P). SPAIN Cantabria, Meron, San Vicente de la Barquera, sand dunes, 30 August 1983, C. Aedo (MA no. 622313). El Tejo, Santander, 18 August 1968, M. R. de la Fuente (MA no. 682797) and dunes, 30 June 1969, M. R. de la Fuente (MA no. 682794). Liencres, Santander, dunes, 18 May 1969, M. R. de la Fuente (MA no. 682798) and 24 May 1969, M. R. de la Fuente (MA no. 682799). Cuchia, Santander, 2 July 1971, M. R. de la Fuente (MA no. 682796). Loredo, Ribamontan al Mar, dunes, 12 June 1993, M. R. de la Fuente (MA no. 682795), and 19 May 1990, C. Aedo (MA no. 622378). Loredo, Santander, Port-duna-arenosa, 7 October 1979, M. R. de la Fuende (MA no. 682825). Watsonia 26: 443-450 (2007) 443 Pollen studies in British Hieracium sect. Alpina (Asteraceae) K. SLADE*' and T. C. G. RICH*? Department of Biodiversity and Systematic Biology, National Museum of Wales, Cardiff CF 10 3NP ABSTRACT A survey of pollen production in British Hieracium sect. Alpina (Asteraceae) taxa has been undertaken using Alexander’s Stain as a test for potential pollen viability. Diploid H. alpinum s.s. from Europe produces abundant, well-formed, stainable pollen. No pollen was observed in British H. alpinum s.s. Potentially viable pollen was observed in 14 of the 39 sect. Alpina taxa investigated, including H. holosericeum (triploid), H. leptodon (tetraploid) and H. pentaploideum (pentaploid), sometimes in abundance. Cultivated plants were more likely to have pollen than wild plants, suggesting pollen production may be under both environmental and genetic control. The involvement of pollen should be considered in theories of evolution of Hieracium sect. Alpina. KEYWORDS: Hieracium alpinum, pollen viability, Alexander’s Stain, Britain. INTRODUCTION As far as it is known, all the British taxa of Hieracium sect. Alpina (Griseb.) Gremli are polyploid, obligate apomicts (Stace ef al. 1997). In Britain, the taxa are triploid (3n = 27), tetraploid (4n = 36), or rarely pentaploid (Sn = 45) (Stace et al. 1995). The apomictic reproductive system is by diplospory (as has been shown for H. alpinum sensu stricto by Skawinska (1962)), in which the embryo is de- rived directly from an unreduced archesporial cell, with no male gamete input either to the embryo, or for formation of the endosperm (i.e. they are not pseudogamous). Thus reproduction in these British apomictic polyploids is essentially clonal, with progeny genetically identical to the mother and to each other. In Europe, both diploids and polyploids occur in sect. Alpina. Chrtek (1997) and Strochova et al. (2002) showed that H. alpinum in the eastern Carpathians is a diploid, sexual, self- incompatible taxon, with apomictic triploid clones elsewhere (including Britain) which did not produce pollen. *!e-mail: Katherine.Slade@museumwales.ac.uk **e_mail: Tim.Rich@museumwales.ac.uk The main theory to explain the evolution of polyploid apomictic groups such as Hieracium sect. Alpina is that they have evolved through hybridisation. Hybridisation may have occurred between two sexual taxa, with the progeny subsequently becoming apomictic and poly- ploid independently. Alternatively, hybrids might have arisen from pollination of a sexual species by an apomictic polyploid whose apomictic genes are then expressed in sub- sequent generations. A combination of both mechanisms may have occurred. Other possible Speciation mechanisms include point mutations, ‘pseudo-sexual’ recombination among different copies of the same chromosomes within the mother, and chromosome structural mutations which simultaneously affect numerous genes and traits (Tyler 2006). Of the British sect. Alpina 34 of the 35 species are endemic, but it is not known where they originated or what their parentages are (Stace et al. 1997). As diploid H. alpinum is currently only known from the Carpathians, and is also the only known diploid in sect. Alpina, it is possible that some of our endemics originated in the Carpathians and then migrated to Britain. For the taxa to then become endemic to Britain, they would need to evolve independ- ently in Britain and/or Europe, and diverge so much that they are recognised as distinct taxa. Alternatively, it is possible that diploid H. alpinum migrated to Britain after the last glacial period, gave rise to some of our endemics and then died out. The British endemics could also have arisen from some closely related sexual taxa, not necessarily diploids, which had migrated to Britain. Stace et al. (1997) investigated the cyto- logical and molecular variation in British sect. Alpina, and found that despite being apomictic, there is genetic variation both between and within taxa. The origin of the variation could be explained in parts by mutations, poly- phyletic origin and relics of sexual hybrid- isations. As the amount of genetic variation 444 KeSEADETS ale CaGeRi@h observed was low, they suggested that the same or similar sexual parents, both diploid and probably tetraploid, gave rise repeatedly to a series of different diploid, triploid, tetraploid and pentaploid hybrids in more than one locality. They suggested that if diploid H. alpinum had been present in Britain, it may have died out due to low seed set due to its requirement for cross-pollination which would have been unpredictable in our suboptimal climate. The low levels of genetic diversity within the sect. Alpina species also suggested recent origins (presumably post-glacial), with some subsequent mutations. As part of work trying to understand evol- ution of the British sect. A/pina endemics, a survey of pollen production was undertaken using Alexander’s Stain (Alexander 1969). Alexander’s Stain stains the outside of the pollen grain greenish-blue and cytoplasm inside red, so well-formed pollen grains with cytoplasm (which are potentially viable) can be picked out quickly. It is a quick method to screen many plants, including historical herbarium specimens, for potential fertility, but does not prove that the grains are viable. It is generally reported that the polyploid Hieracium apomicts produce no pollen or only small amounts of inviable pollen (e.g. Stace er al. 1997), but we have recently observed reasonable quantities of apparently viable pollen in both herbarium specimens and living material. The involvement of pollen in sexual reproduction might help to explain evolution in the genus and the concentrations of endemic species. Hieracium alpinum was investigated in detail due to the potential role of sexual diploids in the origin of the endemics, with a selection of material of the remaining British species of sect. A/pina for comparison. METHODS Pollen samples of British H. alpinum were taken from herbarium specimens in BM, E and NMW, aiming to cover as much of the geo- graphic range as possible. Diploid material from Europe was provided by P. Mraz. Specimens of the remaining British sect. Alpina species were sampled in NMW. The accession numbers of all herbarium sheets examined are listed in Appendix 1. Capitula were examined under a microscope and the ring of anthers removed from florets about to open, or where not available from recently opened florets with emerging styles, using tweezers and a low-power binocular microscope. At least two florets from each specimen were examined, sometimes more. The anthers were placed onto a slide with a drop of Alexander’s Stain, warmed briefly on the hotplate, then broken up with the tweezers to release the pollen, and the tissue ‘stroked’ to free the pollen grains stuck inside the anthers. The preparation was then covered with a cover slip and either placed onto a hotplate to improve the uptake of the stain for at least five minutes or left overnight. The slides were then examined under a high- power compound binocular microscope. The areas on the slide with high numbers of pollen grains were sampled to estimate the percentage of fertile grains. Grains present on the stigmas were ignored as they may have come from another source. Numbers of potentially fertile and infertile grains were recorded, and the percentage fertility calculated. RESULTS Two main types of pollen grains were found. First, spherical grains with the cytoplasm inside which stained bright red and thick papillate cell walls which stained green. The grains could vary in size, sometimes more so in one species than another. These pollen grains were taken as potentially viable. Second, spherical grains staining green only, with no red staining inside (i.e. no cytoplasm). These were sometimes smaller in size than those with cytoplasm, but very few were deformed. These grains were considered infertile. Potential pollen viability in European material of diploid H. alpinum is shown in Table 1. The specimens examined generally produced abundant, well-formed pollen of uniform size which had a high stainability. In contrast, no pollen was found in the 85 British H. alpinum herbarium specimens, apart from in two specimens of doubtful origin or identification. The locations of H. alpinum specimens from which we have examined for pollen are shown in Figure 1. Surprisingly, 14 of the 39 British Hieracium sect. Alpina taxa produced large amounts of pollen (Table 2). Where pollen was present within a taxon, it was not necessarily present in all individuals examined, and it varied in quantity between individuals. Furthermore, potential viable pollen was observed across the three higher ploidy levels (Table 3) including H. holosericeum (triploid), H. Jleptodon (tetraploid) and H. pentaploideum (pentaploid). POLLEN STUDIES IN BRITISH HIERACIUM SECT. ALPINA 445 TABLE 1. POTENTIAL POLLEN VIABILITY IN DIPLOID H. ALPINUM. (TOTAL NUMBER OF POLLEN GRAINS COUNTED ARE GIVEN IN BRACKETS) Origin No. florets examined Potential pollen viability Chornohora Mountains, Ukraine, plant | D, 99% (n = 50) Chornohora Mountains, Ukraine, plant 2 99% (n = 50) Chornohora Mountains, Ukraine, plant 3 41% (n= 86) 99% (n = 50) 96% (n = 50) Rodna Mountains, Romania Bucegi Mountains, Romania NON NWN FIGURE 1. Localities from which no pollen was found in the 85 Hieracium alpinum herbarium specimens examined (@). Other H. alpinum localities from which no material has been examined (O). All records are shown irrespective of date. 446 Ke SEADET Se Cy GaRICEH TABLE 2. POLLEN PRODUCTION IN BRITISH SECT. ALPINA SPECIES. PLOIDY LEVEES ARE TAKEN FROM STACE ET AL. (1995, 1997). TOTAL NUMBER OF POLLEN GRAINS COUNTED ARE GIVEN IN BRACKETS Taxon Ploidy No. plants Potential pollen viability (%) examined H. backhousei F. J. Hanb. x4 2 No pollen H. calenduliflorum Backh. x4 2 No pollen H. calvum P. D. Sell & D. J. Tennant x4 D No pollen H. completum P. D. Sell & C. West x4 3 45% (n= 115), 2 plants no pollen H. deargicola P. D. Sell & D. J. Tennant x4 i No pollen H. einichense P. D. Sell & D. J. Tennant x4 ] No pollen H. eximium Backh. x4 3 No pollen H. eximium Backh. f. eximium x4 2, No pollen H. eximium Backh. f. tenellum (Backh.) P. D. Sell x4 3 No pollen & C. West H. globosiflorum Pugs}. x3 3 No pollen H. graniticola W. R. Linton x3 2 No pollen H. grovesii Pugsl. x4 3 11% (n= 63), 13% (n= 111), 6% . (n = 63) H. hanburyi Pugsl. f. hanburyi x4 12 No pollen H. hanburyi Pugsl. f. atraticeps (Pugsl.) P. D. x4 yD, 4% (n = 22), | plant no pollen Sell & D. J. Tennant H. hanburyi Pugsl. f. pusillum P. D. Sell & D. J. x4 ] No pollen Tennant H. holosericeum Backh. x3 10 24% (n = 50), 8% (n = 50), 8 plants no pollen H. insigne Backh. f. insigne x4 ] Underdeveloped pollen H. insigne Backh. f. celsum P. D. Sell & C. West = x3 or x4 5 35% (n = 47), 12% (n = 68), 100% (n = 31), 34% (n = 89), 1 plant no pollen H. kennethii P. D. Sell & D. J. Tennant x4 I 8% (n = 23) H. larigense (Pugsl.) P. D. Sell & C. West unknown ] No pollen H. leptodon P. D. Sell & D. J. Tennant x4 6 64% (n = 62), 46% (n = 90), 33% (n = 24), 3 plants no pollen H. macrocarpum Pugsl. x4, x4+] 3 86% (n = 50), 2 plants no pollen H. marginatum P. D. Sell & C. West x4 2 94% (n = 72), 61% (n = 103) H. memorabile P. D. Sell & C. West x4 = No pollen H. milesii P. D. Sell & C. West x4 3 No pollen H. mundum P. D. Sell & C. West x4 2 29% (n= 106), 1 plant no pollen H. notabile P. D. Sell & C. West x4 D, No pollen H. optimum P. D. Sell & C. West unknown 2 No pollen H. pensum P. D. Sell & C. West x4 ] 2570») H. pentaploideum P. D. Sell & D. J. Tennant x5 2 68% (n = 69), 100% (n = 50) H. perscitum P. D. Sell & C. West x4 Z 24% (n= 148), 1 plant no pollen H. probum P. D. Sell & C. West x4 5 No pollen H. pseudocurvatum (Zahn) Pugsl. x3 ] No pollen H. pseudopetiolatum (Zahn) Roffey x3 3 12% (n = 26), two plants no pollen H. subglobosum P. D. Sell & C. West x3 4 1% (n = 50), 3 plants no pollen H. subgracilentipes (Zahn) Roffey x4 1 No pollen H. subtenuifrons P. D. Sell & D. J. Tennant x3 or x4 3 No pollen H. tenuifrons P. D.S ell & C. West x4 4 36% (n= 112), 68% (n = 105), 2 plants no pollen POLLEN STUDIES IN BRITISH HIERACIUM SECT. ALPINA 447 TABLE 3. NUMBER OF TAXA IN EACH PLOIDY LEVEL AND POLLEN PRODUCTION IN BRITISH SECT. ALPINA SPECIES Ploidy Total no. taxa Pollen absent Significant pollen production x3 6 + 2 x3 or x4 2 ] l x4* DT 16 11 x5 1 0 1 Total 36 21 IS *includes H. macrocarpum aneuploid It is unclear why pollen should be present in some individuals of a taxon, but absent in others. We noticed that cultivated plants were more likely to have pollen than plants collected nomic wild: For imstance, in 4. subglobosum, the cultivated specimen had masses of pollen (although with only 1% viability) but the wild-collected plants had none. Excluding H. alpinum, 13 of the 28 cultivated specimens of sect. Alpina species had pollen (46%), compared to only 14 of the 87 wild- collected specimens (24%); this is significantly different (x7, p<0-001). DISCUSSION The key result is the observation that although no pollen was observed in British H. alpinum, significant amounts of potentially viable pollen were observed in over a third of the other sect. Alpina taxa. The stain does not prove that pollen is viable, which must be tested by germinating fresh pollen from cultivated plants, but the presence of pollen nonetheless suggests the possibility that it might have a role in evolution of the endemics. Our survey of pollen on _ herbarium specimens shows there is no evidence that diploid H. alpinum occurs in Britain. Although there are only counts of three plants from two localities published (Stace ef al. 1995), it is presumably uniformly triploid. Strochova ef al. (2002) showed allozyme and RAPD variation occurred within and between populations of the triploid cytotype of H. alpinum, and the geographic and genetic distances were found to be correlated, indicating isolation by distance. They interpreted the gradient of genetic variation observed from the variable diploids in the Eastern Carpathians to the uniform triploids elsewhere as resulting from the post-glacial recolonization of Europe from glacial refugia in the Carpathians. Diploids may therefore never have been present in Britain. If diploid H. alpinum has never been present in Britain, did the endemic taxa evolve here by hybridisation or migrate? The endemics are unlikely to have evolved from hybridisation with diploids in Britain as the only sexual, diploid species of Hieracium present is H. umbellatum L., which is so morphologically different that it cannot have been involved. The occurrence of pollen in polyploids suggests that the endemics could have evolved by hybrid- isation between some endemic taxa if the polyploids are partly sexual: this would require both fertile pollen and a diplospory apomictic reproductive system that is not absolute (i.e. occasional fusion of gametes may occur). Detection of such hybrids within sect. Alpina would be difficult as the species are generally very similar, differing in few characters and showing a reticulate pattern of morphological variation. Hieracium hybrids have only very rarely been observed in the wild (Mraz ef al. 2003) or cultivation (Pugsley 1948). Migration of at least some parental taxa which sub- sequently became endemic in addition to H. alpinum is still required under this model. Hybridisation between polyploids could explain how the endemic H. pentaploideum arose on An Teallach, Wester Ross (v.c. 105) from the other species recorded on _ the mountain. It could have resulted from a cross of an unreduced triploid gamete from H. alpinum or H. holosericeum with a reduced diploid gamete from a tetraploid such as H. kennethii, H. marginatum or H. perscitum. Similarly, Sell & Murrell (2006) note how species within the artificial group sect. Subalpina may have arisen as hybrids between sect. Alpina species and species within other sections such as sect. Cerinthoidea, sect. Oreadea and sect. Vulgata. Given that 42 of the 45 sect. Swbalpina species are endemic, this 448 hybridisation is again likely to have happened between polyploids in Britain. If the British sect. Alpina endemics did migrate from their place of origin in Europe, it K. SLADE & T. C. G. RICH and the higher pollen production in cultivated specimens suggests that pollen production may be under environmental as well as genetic control. is surprising that only H. alpinum still occurs there; it is difficult to explain how the 34 other species followed the same or very similar migration pattern. The involvement of pollen should be cons- idered in theories of evolution of Hieracium sect. Alpina, and experimental work to test this hypothesis is required. The variation in pollen production between specimens within a species ACKNOWLEDGMENTS We would like to thank Patrik Mraz for the samples of European diploid H. alpinum and the Keepers of Natural History Museum and Royal Botanic Gardens Edinburgh for loan of Hieracium alpinum. REFERENCES ALEXANDER, M. P. (1969). Differential staining of aborted and non-aborted pollen. Stain Technology 44: 117— W222, CHRTEK, J. JNR (1997). Taxonomy of the Hieracium alpinum group in the Sudeten Mountains, the West and the Ukrainian East Carpathians. Folia Geobot. Phytotax., Praha 32: 69-97. MRAZ, P., CHRTEK, J. & PLACKOVA, I. (2003). First record on recent natural hybridisation in the genus Hieracium s. str. In, Hieracium workshop, Krivoklat, Czech Republic, 28 May — I June, 2003. Institute of Botany, AS CR & Protected Landscape area “Krivoklatsko” [sine pag. | PUGSLEY, H. W. (1948). A prodromus of the British Hieracia. Journal of the Linnean Society (Botany) 54: 1— 356. SHI, Y., GORNALL, R. J., DRAPER, J. & STACE, C. A. (1996). Intraspecific molecular variation in Hieracium sect. A/pina (Asteraceae), an apomictic group. Folia Geobotanica & Phytotaxonomica 31: 305-313. SKAWINSKA, R. (1962). Apomixis in Hieracium alpinum L. Acta Biologica Cracoviensa, Series: Botanica 5: 89-96. STACE, C. A., GORNALL, R. J., SQUIRRELL, J. & SHI, Y. (1995). Chromosome numbers in Hieracium L. section Alpina (Fries) F. N. Williams. Watsonia 20: 367-377. STACE, C. A., GORNALL, R. J. & SHI, Y. (1997) . Cytological and molecular variation in apomictic Hieracium sect. Alpina. Opera Botanica 132: 39-51. ‘ STORCHOVA, H., CHRTEK, J. JNR, BARTISH, I. V., TETERA, M., KIRSCHNER, J. & STEPANIK, J. (2002). Genetic variation in agamospermous taxa of Hieracium sect. Alpina (Compositae) in the Tatry Mts. (Slovakia). Plant Systematics and Evolution 235: 1-17. TYLER, T. (2006). Patterns of morphometric variation and a new supraspecific classification of apomictic taxa of Hieracium (Asteraceae) from Denmark and southern Sweden. Plant Systematics and Evolution 261: 39-88. (Accepted April 2007) POLLEN STUDIES IN BRITISH HIERACIUM SECT. ALPINA 449 APPENDIX 1. LIST OF ACCESSION NUMBERS OF HERBARIUM SHEETS OF BRITISH HIERACIUM SECT. ALPINA EXAMINED. SPECIMENS PREFIXED BM ARE FROM THE NATURAL HISTORY MUSEUM, LONDON, AND PREFIXED E FROM ROYAL BOTANIC GARDENS, EDINBURGH. ALL OTHER SHEETS ARE IN NMW H. alpinum: 25.148.516, 25.149.5156, 27.72.1260, 28.131.4188, 28.131.4242, 49.29.5538, 85.40.2659, V.2005.1.233, BM000916035, BM000916037, BM000916049, BM000916050, BM000916052, BM000916053, BM000916054, BM000916055, BM000916056, BM000916057, BM000916058, BM000916062, BM000916063, BM000916064, BM000916065, BM000916066, BM000916067, BM000916068, BM000916069, BM000916070, BM000916071, BM000916072, BM000916073, BM000916074, BM000916075, BM000916076, BM000916077, BM000916078, BM000916082, BM000916083, BM000916084, BM000916085, BM000916106, BM000916086, BM000916087, BM000916088, BM000916089, BM000916090, BM000916091, BM000916092, BM000916093, BM000916094, BM000916095, BM000916096, BM000916097, BM000916098, BM000916099, BM000916100, BM000916101, BM000916102, BM000916103, BM000916104, BM000916105, BM000916107, BM000916108, E0003 1564, E00031565, E00031566, E00031567, E00031568, E00031569, E00031571, E00031572, E00031573, E00031574, E00031575, E0003 1576, E00031577, E00031578, E00031580, E00031581, E00031650, E0003 1663, E00048447, E00048602, E00171460, E00209980, E00209981. H. backhousei: V.2000.008.6, V87.58.3081. H. calenduliflorum: 27.72.1265, 28.131.4227. H. calvum: V.2000.008.28, V.2000.008.30. H. completum: 25.149.5178, 25.149.5195, V87.58.3080. H. deargicola: V.2000.008.58. H. einichense: V.2000.008.59. H.. eximium: 27.72.1268, 28.131.4206, 28.131.4219, 26.553.574, 28.131.4215a. H. eximium f. tenellum: 27.72.1269, 28.131.4217. H. globosiflorum: 20.347 .29, 27.72.1267, 28.131.4244. H. graniticola: 28.131.4239, V.2000.008.10. H. grovesii: V.2000.008.12, V.2000.008.14, V.2000.008.15. nemnanouniz N 2005.1.234, 25.149.5176, 25.149.5188, 25.149.5192, 25.49.5193, 27.72.1273, 28.131.4303, 28.131.4304, 28.131.4306, 28.131.4307, 28.603.23, V87.58.3084. H. hanburyi f. atraticeps: 85.40.2660, V.2000.008.5. FH. hanburyi f. pusillum: V.2000.008.16. Pmmnolosericcum: 202, 26.553.5)3, 27.12. 1262a, 28. 130-4192, 28.131-4194, 228.131.4195, 28.131.4196, 28.603.69, 49.29.5536, V87.58.3093. H. insigne f. celsum: V.2000.008.34, V.2000.008.35, V.2000.008.36, V.2000.008.37, V.2000.008.38. H. insigne f. insigne: V.2000.008.39. H. kennethii: V.2000.008.40. H. larigense: V.2000.008.41. H. leptodon: 28.131.4211, 28.131.4212, 28.131.4214, 28.131.4229, V.2000.008.42, V.2000.008.43. Hf. macrocarpum: 28.131.4246, V.2000.008.17, V87.58.3095. H.. marginatum: V.2000.008.61, V.2000.008.19. H. memorabile: 28.131.4257, 28.131.4258, 28.131.4260, V.2000.008.20. H. milesii: 28.131.4237, 49.29.5534, V.2000.004.27. H. mundum: 28.131.4259, V.2000.008.51. H. notabile: 28.131.4225, V.2000.008.21. H. optimum: 28.131.4213, V.2000.008.52. 7. pensum: V .2000.008.53. H. pentaploideum: V.2000.008.60, V.2000.008.60. 450 Ke SEADE ely 2G, RICH H. perscitum: 28.131.4248, V.2000.008.54. H. probum: 25.149.5179, 28.131.4238, 28.131.4253, 28.131.4319, V.2000.008.55. H. pseudocurvatum: 28.131.4255. H. pseudopetiolatum: V.2000.008.22, V87.58.3078. H. subglobosum: 25.149.5173, 25.149.5177, 28.131.4247, V.2000.008.56. H. subgracilentipes: V.2000.008.24. H. subtenuifrons: V.2000.008.57, V.2000.008.63, V.2000.008.63. H. tenuifrons: 28.131.4197a, 25.149.5169, 28.131.4209, 28.131.4231. Watsonia 26: 451-461 (2007) 45] Hieracium portlandicum (Asteraceae), a new endemic hawkweed from the Isle of Portland, England related to Hieracium leyanum T. C. G. RICH'* Department of Biodiversity and Systematic Biology, National Museum of Wales, Cardiff CF 10 3NP B. EDWARDS Dorset Environmental Records Centre, Library Headquarters, Colliton Park, Dorchester DTI LXJ and D. A. PEARMAN** Algiers, Feock, Truro, Cornwall TR3 6RA ABSTRACT A new species of hawkweed is described. Hieracium portlandicum differs from H. /evanum, within which it was previously included, in being less hairy, and in having narrower, acute, light greyish-green, less hairy involucral bracts. It is a triploid apomict. Hieracium leyanum is a tetraploid apomict. Hieracium port- landicum is endemic to the Isle of Portland, Dorset, England, where it occurs on Portland Limestone rocks, and on soils derived from it. 103 plants are currently known from two sites, and a comparison against historical data indicates that it is declining. It is ‘Endangered’ under the IUCN Threat Criteria. Some distribution data are also given for H. leyanum which is ‘Vulnerable’ under the IUCN Threat Criteria. A lectotype for H. pollinarium var. platyphyllum on which H. leyanum is based 1s designated from Wales. KEYWORDS: Compositae, chromosome number, lectotype, pollen viability, reproductive biology. INTRODUCTION The identity of a Hieracium species of the Isle of Portland, Dorset (v.c. 9) has been uncertain since it was first found by W. C. Medlycott in iomomelbcyancr al. (1908) referred! it to 7 leyanum (Zahn) Roffey (under the synonym H. platyphyllum), and noted that the Portland record was a curious extension of range for what was otherwise an endemic species of *e-mail: tim.rich@museumwales.ac.uk *e-mail: dpearman4@aol.com South Wales and the adjacent English counties. E. F. Linton (in Ley et a/. 1908) also noted that the Portland plant differed from H. /Jeyanum in characters which might have been due to climate and situation, and suggested it would be worthwhile cultivating the two forms side by side to test the identification. Linton’s differences between the taxa noted on a herbarium sheet in BM, updated with modern terminology and with missing words added in square brackets, are as follows: H. leyanum (as H. platyphyllum): Leaves with stellate hairs and scattered [simple eglandular] hairs beneath and on the margins; capitula rather large, thick truncate-based; involucral bracts big, broad, with long [simple eglandular] hairs and sparse stellate hairs. Portland plant: Leaves hairy on both surfaces; capitula medium (not large); involucral bracts [small, narrow], with short, black-based [simple eglandular] hairs and stellate hairs; shortish glands very few on base of involucral bracts and peduncles. Pugsley (1948) noted that the Portland plant differed materially from H. /eyanum in having much narrower involucral bracts and deserved further investigation. Sell & West (1968) agreed it differed slightly but not enough to merit specific rank, and noted that the plant had not been seen in Portland ‘in recent years’. Bowen 452 T. C. G. RICH, B. EDWARDS & D. A. PEARMAN (2000) listed it from four localities, including two recent ones, and noted that his use of the name H. /eyanum was provisional, and his card index (currently held by D.A.P.) gives an additional site. Sell & Murrell (2006) retain it within H. /eyanum and also suggest that the Portland plants are a good match for H. rubicundiforme (Zahn) Roffey. In 20035 C5 Ge Rich™ began collating: information on H. /eyanum as part of a project on critical Welsh taxa and, intrigued by the isolated Portland locality, asked B. Edwards and D. A. Pearman to collect Portland seed to grow for comparison with Brecon material. Seed was duly collected, and following cultivation of plants beside each other, as suggested by Linton (Ley ef al. 1908), the Portland plant is now considered to merit recognition as a species in its own right. METHODS MORPHOLOGY Seeds were collected from the one fruiting plant seen at West Weare, Portland on 20 July 2003 by B. Edwards and D. A. Pearman, and from several plants in each of Craig Cerrig- gleisiad, Cwm Tarrell and Darren Fach in Brecon im’) 2003) by, ©) GRichScedsawenre sown in damp John Innes no. 1 potting compost in a greenhouse and the resulting plants were grown on. Most seeds of the Brecon plants germinated in autumn 2003. Seedlings remained green during the winter, and they first flowered in June 2004. Three seeds of the Portland plant germinated in the spring of 2004 and flowered about three weeks after the Brecon plants. In 2005, one Portland plant flowered about five weeks after the Brecon plants (the other plant did not flower). The plants were compared side by side, and one specimen from each site was pressed (NMW). Plants were also examined in the field, and herbarium material was examined in BEL, BIRM, BM, CGE, DOR, E, LIV, MANCH, NMW and RNG. Material of C. West in MNE has not been examined. Terminology of hair lengths follows Sell & Murrell (2006). CHROMOSOME COUNTS Tips of actively growing roots were collected mid-morning and pre-treated in a 8-hydroxyl- quiniline for 20-24 hours in the fridge. They were then fixed in a 3:1 mixture of absolute ethanol: glacial acetic acid and stored until needed in the fridge. Root tips were then hydrolysed in concentrated 5N HCl for 10 minutes at room temperature, then stored in 70% industrial methylated spirits. Meristems were dissected out in 45% acetic acid, and squashed between a microscope slide and coverslip in 0-1% aceto-orcein stain with brief flame treatment. REPRODUCTIVE BIOLOGY Two florets were taken from each herbarium sheet to assess potential pollen viability (Cleal & Rich 2004). Anther rings were removed from florets with tweezers under a low-power binocular microscope, and placed onto a slide with a drop of Alexander’s Stain (Alexander 1969), warmed briefly on the hotplate, then broken up with the tweezers to release the pollen. The preparation was then covered with a cover slip and replaced on the hotplate to improve the uptake of the stain. The slides were examined under a high-power compound binocular microscope for areas of dense pollen grains. Potentially viable grains were counted as those which were spherical with thick papillate cell walls which stained green and had cytoplasm inside which stained uniformly bright red. Deformed grains (rare) or those staining green only with very little or no red staining inside (i.e. with little or no cytoplasm) were considered infertile. To test for apomixis, young but well- developed capitula were cut with scissors at about their middle to excise the anthers and styles, before the florets opened, and seed set recorded. Control capitula on adjacent ped- uncles were left for comparison. FIELD SURVEYS Historical and recent records from Portland (Appendix 1) were used to plan the field survey in 2005. These indicated that H. /evanum sensu lato had been recorded in three areas on Portland: West Weare, Verne plateau, and more extensively down the east side from East Weare to Church Ope Cove. On 22 June 2005 these sites were visited by B. Edwards and T. Rich, and data were collected on habitat, population sizes, soils and associated plant species. Soil pH was measured with a pHep2 Hanna pocket-sized pH meter in a 50:50 mixture with distilled water on soil samples collected from around the roots. Details of H. Jevanum were collected during field surveys of other rare Welsh hawkweeds between 2000 and 2003 by T. Rich, with more detailed surveys being carried out specifically for it in 2004 and 2005. HIERACIUM PORTLANDICUM 453 RESULTS MORPHOLOGY Cultivated plants from Portland and from Brecon were vegetatively similar in leaf shape and size, with the exception that the Portland plants lacked the glaucous bloom of Brecon H. leyanum and had more simple eglandular hairs on the stems, petioles and leaves. Both taxa had hairs on both sides of the leaves, but the Portland plants were more densely hairy. The youngest leaves of both species tended to be glabrescent on the upper surface. In flower, all plants had capitula with at least some of the characteristic stylose florets (i.e. the ligules do not fully develop and remain tubular with exserted styles), though more of the florets were stylose in the Brecon plants (this can however be quite variable even within a population, e.g. as observed at Cwm Tarrell in 2004, or Dyffryn Crawnon in 2005). The Brecon plants had capitula with broader bases and slightly larger ligules — to 25 mm long — compared to more rounded bases and ligules to 21 mm in the Portland plant, but the variation in extent of development of stylose florets made systematic comparison difficult. The involucral bracts were clearly different; those of the Portland plants were narrower (as noted by Pugsley 1948), acute, lighter greyish-green with fairly numerous, short (to 0-4 mm), black- based glandular hairs, a few, scattered, long simple eglandular hairs and fairly numerous, appressed stellate hairs, and those of the Brecon plants were broader, obtuse, darker greyish-green, with dense, short to medium length (to 0-6 mm), black glandular hairs, some scattered, long simple eglandular hairs and dense, appressed stellate hairs. The main differences between herbarium specimens from Portland and from Brecon- Hereford were largely as set out by Linton on the BM herbarium sheet (cf. above) and as observed in the cultivated material. The density of stellate hairs on the leaves of Brecon- Hereford H. leyanum was variable, ranging from virtually absent in some plants to sparse in others; most specimens of the Portland plants lacked stellate hairs on the leaves, but a few had very sparse stellate hairs near the edges. The leaves of the Portland plants were noticeably more hairy on the upper surface than those of Brecon-Hereford H. /eyanum, some plants of the latter being + glabrous on the upper surface, and the leaves of both taxa were hairy below. Overall, the wild material was quite variable in hairiness at least at West Weare; sometimes plants were moderately hairy, sometimes they had dense, long, villose hairs. Portland plants were consistently more hairy than Brecon-Hereford plants. Long simple eglandular hairs were much more frequent on the involucral bracts of Brecon- Hereford material than in Portland material. The mean width of the long middle involucral bracts measured at about their middle was 1-54 mim 2 0-06 se. (n= 21, range 1—1-8(-2:4) mm) for Brecon-Hereford H. leyanum, and 0:96 mm + 0-12 s.e. (n = 25, range 0-7—1-3 mm) for the Portland plants; a t-test showed that these widths were significantly different (p<0-001). Comparison of the herbarium material with cultivated material of the same origin showed consistent differences between the Portland and Brecon plants, indicating that the cultivated plants are representative of the character differences in wild plants, though there was overall more variation in hairiness in the wild. The glandulosity of the involucral bracts were more difficult to determine, probably due to drying and storage of the plants, and the differences in leaf coloration between fresh plants were not apparent in herbarium material. In the field, the most immediately apparent gross differences between Portland and Brecon plants were the differences in size, basal rosette leaves and development of the stylose capitula. Most of the Portland plants were small (up to 15 cm high) and slender (Barrett (1912) quotes measurements taken by Mr Fulleylove in 1911 from the east side as ranging from 7:5—45 cm, and averaging 27 cm), compared to the bigger Brecon plants — usually at least 35—70 cm tall, with two or three robust stems. The Brecon plants had noticeably glaucous, larger and relatively broader basal rosette leaves with few or no hairs on the upper surface, whilst the Portland plants had basal rosette leaves green, smaller and narrower, and hairy on both surfaces. Most capitula of the Portland plants seen in 2005 were normal and well-developed, whilst most of the Brecon plants had most capitula stylose, with a few normally dev- eloped capitula. CHROMOSOME COUNTS One count of 2n = 27 (triploid) was obtained for each of two different plants grown from the seed collected from one plant at West Weare, Portland in 2003. Three counts of 2n = 36 (tetraploid) and one of 2n = c. 37 were obtained 454 T. C. G. RICH, B. EDWARDS & D. A. PEARMAN TABLE 1. POTENTIAL POLLEN VIABILITY IN H. LEYANUM S.L. ASSESSED USING ALEXANDER’S STAIN Locality Accession no. Date % viable* Black Mountains, Brecon NMW.25.149.5446b 1898 Dp SP) Black Mountains, Brecon NMW.25.149.5446 1898 0, 6 Black Mountains, Brecon NMW.25.149.5446c 1898 6,22 Craig Cerrig-gleisiad, Brecon NMW.V.2003.1.9 2003 8, 14 Craig Cerrig-gleisiad, Brecon, cultivated NMW.V.2004.3.184 2004 4,48 Darren Fach, Brecon NMW.V.2000.9.710 2000 66, 68 Darren Fach, Brecon, cultivated NMW.V.2004.3.183 2004 10, 40 Storey Arms, Brecon NMW.V.2004.3.180 2004 40, 56 Average 28:9'\G Owe») Church Ope Cove, Portland BM000792509 1931 onl2 Portland NMW.85.58.3164 1907 12, kG Portland BM000792507 1907 IO, 3 Portland BM000792504 1907 10, 24 West Weare, Portland, cultivated NMW.V.2004.3.182 2004 6, 8 . Average 13-6 (4 2:65 s.e.) * Two florets examined for each accession for plants grown from seed from Craig Cerrig- gleisiad, Brecon and one count of 2n = 36 (tetraploid) for plants grown from seed from Darren Fach, Brecon. REPRODUCTIVE BIOLOGY Neither taxon produced much pollen; potential pollen viability was generally more variable in Brecon-Hereford H. Jeyanum than in_ the Portland plants (Table 1). Although Brecon plants on average had about twice as much viable pollen as the Portland plants, a t-test showed no significant difference in potential pollen viability between the two taxa (P>0-06). One excised capitulum on each of two plants from Brecon and on one plant from Portland, and all control capitula, set seed, indicating that reproduction is apomictic in both Brecon and Portland plants, as it is for most other British Hieracia (vouchers in NMW). Pollen may therefore not be involved in seed production. FIELD SURVEY 2005 At West Weare, Portland, 98 plants were found in block boulder scree in five scattered subpopulations of varying size (2, 3, 22, 35 and 36 plants). On the east side of the island, where there were probably once extensive popul- ations, only five widely-separated plants were found between Church Ope Cove and Durdle Pier (population sizes 1, 1, 1 and 2 plants) on the side or tops of larger boulders on the more prominent ridges in the boulder field. Both of these areas were very difficult to search, due to the nature of the terrain, and there is little doubt that more plants are present, but the decline from being ‘plentiful’ between Durdle Pier and Church Ope Cove (Barrett 1912) to ‘rare’ in 2005 is real. No plants were found at East Weare or Verne, most of the latter area either having been quarried or modified since it was recorded in 1925. Full details of the popul- ations have been deposited at the Dorset Environmental Records Centre, English Nature and the B. S. B. I. Threatened Plants Database. The Portland plants varied in size from small plants c. 10 cm tall with no or one stem leaf and two or three capitula, on the tops and sides of boulders, to larger plants 30-40 cm tall with one or rarely two stems leaves and many stems and capitula, on the deeper soils between boulders or in sheltered crevices. The plants were mostly in full flower at the time of survey, and one or two capitula had already set seed (flowering has been observed continuing until at least mid-July in previous years). Of the 103 plants found, 92 were flowering and 11 were vegetative, and the remains of some dead stems from 2004 showed that many plants, but not all, had flowered in successive years. Examination of the whorls of old leaf scars on the rootstock and stems below the current year’s rosette of one plant indicated that it was at least 15 years old. Regeneration was noted in two of the sub-populations at West Weare, but HIERACIUM PORTLANDICUM not in any of the other sub-populations. The block boulder screes and rocks are relatively unstable as the underlying Jurassic clays slip and move, and the populations probably are somewhat dynamic within this area. The Portland plants were usually either growing directly on boulders of Portland Limestone, rooting into tiny cracks or crevices, or between boulders or in crevices in fine grit derived from the limestone. Two soil samples collected from West Weare had pH values of 6:7 and 6-8. Plants growing directly on rocks with virtually no soil are extremely drought- prone, and sea mist may be important in providing significant water during the summer. The current altitudinal range is c. 10-40 m, though historically the record from Verne plateau would have been at c. 100 m altitude. The vegetation on the rocks or boulder scree was typically very open (from less than 1% to 10% cover) and varied in aspect and slope. It could not be equated to any recognisable plant community. The associated species were various combinations of Asplenium ruta- muraria L., Buddleja davidii Franch., Centranthus ruber (L.) DC., Cymbalaria muralis P. Gaertn., B. Mey. & Scherb., Festuca rubra L., Geranium robertianum L., Hedera helix L., Inula conyzae (Griess.) Meikle, Ligustrum vulgare L., Lonicera periclymenum L., Lotus corniculatus L., Mycelis muralis (L.) Dumort., Pilosella officinarum F. W. Schultz & Sch. Bip., Plantago lanceolata L., Rubia peregrina L., Sonchus oleraceus L., Viburnum lantana L. and rarely Euphorbia portlandica L. Historical and recent records of Hieracium leyanum sensu stricto are listed in Appendix 2. It was refound in most localities and it could still occur at Red Daren and Fan y Big where there are large areas of inaccessible, crumbling rocks, but is unlikely to be still present at Craig y Cilau. It typically occurs in scrub or open rocks on Carboniferous Limestone or Old Red Sandstone, often in small, restricted populations. DISCUSSION To ascertain whether a new name ts needed for the Portland plant, it is first necessary to lectotypify H. leyanum (Zahn) Roffey. The name Hieracium ley[i|anum (Zahn) Roffey was based on H. sagittatum Lindb. ex Stenstr. subsp. /eyianum Zahn. This was a new name for the plant originally described as H. pollinarium F. J. Hanb. var. platyphyllum Ley (=H. platyphyllum (Ley) W. R. Linton) (Zahn 455 1921-1923), which was described from South Wales and Herefordshire by Ley (1898). A lectotype for H. pollinarium F. J. Hanb. var. platyphyllum is hereby designated as the sheet in herb. A. Ley (currently in CGE) collected by him and labelled ‘Taren ‘r Esgob, Breconshire, 12 July 1897’, with the corrected proofs of Ley’s (1898) Journal of Botany note attached. Thus H. leyanum is based on Welsh material. The Portland plants differ consistently from Brecon-Hereford H. Jevanum in morphology of the leaves and involucre, and in chromosome number. They merit recognition as a separate taxon, which for apomictic Hieracium 1s usually most appropriate at species rank: Hieracium portlandicum T. C. G. Rich sp. nov. HOLOTYPUS: on boulder in block boulder scree, West Weare, Portland, Dorset (v.c. 9), England, SY684730, 22 June 2005, B. Edwards & T. C. G. Rich, NMW (accession number V.2005.1.1). Ab Hieracio leyano foliis triste viridibus, in parte superiore pilis simplicibus eglandulosis satis numerosis vestitis, et involucri squamis angustis (circa 0-7—1-:3 mm latis) acutis dilute cinereoviridibus, pilis stellatis appressis satis numerosis pilisque glanduliferis brevibus nigris satis numerosis vestitis, differt. Differs from H. /eyanum in having dull green leaves with fairly numerous simple eglandular hairs above, and narrow (c. 0:7—1-3 mm wide), acute, light greyish-green involucral bracts with fairly numerous appressed stellate hairs and fairly numerous short black glandular hairs. Vernacular name: Portland Hawkweed. Herbaceous, polycarpic, phyllopodous hemi- cryptophyte with branched rootstock and 1-6 stems, stems 9-45 cm high, with sparse to dense, medium length to long (to 3 mm), white simple eglandular hairs throughout and dense stellate hairs above. Basal rosette leaves to 8(— 12) cm long, dull green on the upper surface with minor reddish-purple tints, green or flushed reddish-purplish below, the petioles to 2 cm, winged, tvillous with medium length to long simple eglandular hairs and occasionally sparse stellate hairs, the lamina 3—6(—8) cm x 2—4(—5) cm, broadly elliptic or ovate, truncate to rounded at the base, obtuse-mucronate to acute or acuminate at the apex, the margins sparsely but sharply denticulate towards the 456 base with forward-pointing teeth and remotely glandular-denticulate to entire towards the apex and with dense, ciliate, medium length simple eglandular hairs, moderately hairy with curved, medium length simple eglandular hairs on the upper and lower surfaces, especially on the lower midrib, sometimes densely so, and often more hairy below, the youngest leaves more or less glabrous on the upper surface. Stem leaves usually O—1(—2), similar to basal leaves but usually lanceolate and acuminate at the apex and more sharply toothed, often with much smaller, linear-lanceolate or filamentous bracts at the base of the synflorescence branches (when present). Synflorescence racemose- corymbose with (1—)2—9(-17) capitula, with long, ascending to spreading branches and peduncles, with dense stellate hairs and scattered short, black-based simple eglandular hairs and very short, dark glandular hairs above. Capitula medium-sized c. 35-45 mm in diameter, rounded to truncate at the base. Involucral bracts connivent in bud, in flower 10-12 mm long (11-13 mm to base of involucre), light greyish-green with paler margins, narrow (middle involucral bracts 0-7— 1:3 mm wide at middle), linear-triangular, acute, with fairly numerous, appressed stellate hairs especially on the margins, and with fairly numerous short, black glandular hairs and very few to few, short to medium Inegth, black- based simple eglandular hairs. Ligules to 21 mm (or to 26 mm to base of involucre) but central florets often discoid and shorter, mid- yellow, glabrous at the tips, the tips often variably cut into linear-oblong lobes. Styles discoloured with greyish-black papillae and stigmas. Margins of receptacle pits weakly dentate. Seeds 3-0—-3:8 mm, purplish-black. Flowering time: June—July. Hieracium portlandicum is endemic to the Isle of Portland, Dorset (v.c. 9), England, where it has been recorded in at least three localities (Figure 1). The population survey in 2005 revealed a total of 103 plants, and it has clearly declined on the east side of Portland during the last 100 years, suggesting that it is best treated as ‘Endangered’ under the IUCN Threat Criteria (IUCN 2001). The main threats are currently development of scrub, and especially the spread of alien Cotoneaster species and possibly Hedera helix over its rocky outcrops near Church Ope Cove. The other alien species associated with H. port- landicum do not currently pose a significant threat. No other species of Hieracium are recorded from the Isle of Portland. ; T. C. G. RICH, B. EDWARDS & D. A. PEARMAN Hieracium portlandicum is illustrated in Figure 2, which also shows involucral bracts of H. leyanum for comparison. There is an excellent photograph of H. portlandicum in Bowen (2000, Plate 37). Pugsley (1948) and Sell & Murrell (2006) give descriptions of H. leyanum. Sell & West (1968) noted that H. leyanum, H. angustisquamum (Pugsley) Pugsley and H. holophyllum W. R. Linton had many characters in common with H. caledon- icum F. J. Hanbury, which could be the parent from which they were derived; this is also likely to be the case with H. portlandicum. Hieracium caledonicum, which occurs in northern Scotland, differs from H. port- landicum in having smaller capitula with broader, obtuse involucralebractssee holophyllum differs in having obtuse involucral bracts and yellow styles, and H. angustis- quamum differs in having usually three stem leaves and obtuse involucral bracts. Sell & Murrell (2006) suggested that the Portland plants were a good match for H. rubicundi- forme (formerly H. rubicundum F. Hanb.), but they differ in having green, not deep bluish- green, leaves which are hairy on both sides and in the acute, not obtuse involucral bracts. No chromosome counts are available for these other related taxa. Pugsley (1948) placed Hieracium leyanum in Section Vulgata (Griseb.) Willk. & Lange, but Sell & West (1968) placed it in Section Oreadea (Fr.) Arv.-Touv.; Hieracium portlandicum is best included with H. /eyanum in the latter. Hieracium leyanum sensu stricto has been recorded from 12 sites in four vice-counties (Fig. 3). It has been seen between 2003 and 2005 in eight sites in four hectads, all in v.c. 42 Brecon, with a total population size of at least 800 plants (Appendix 2). It was not refound in Monmouthshire or Herefordshire. It qualifies for listing in the current British vascular plant red data book (Wigginton 1999), and it is best classified as ‘Vulnerable’ under the [UCN Threat Criteria until comprehensive surveys can be carried out (IUCN 2001). ACKNOWLEDGMENTS We would like to thank the Keepers of the Herbaria cited, and John Bailey, Helen Cleal, Jenny Cripps, Trevor Evans, Lady Rosemary FitzGerald, Peta Hayes, Stephen Jury, Alex Lockton, David McCosh, Gina Murrell, Philip Oswald, Mike Porter, Chris Preston and Peter Sell for their help. HIERACIUM PORTLANDICUM 457 Nh eZ * Fortuneswell © Verne Easton y : , 5 & au é a Lt A ys Church Ope Cove FIGURE 1. Distribution of Hieracium portlandicum on Portland, Dorset. @, 2005. Grey shaded, approximate areas where it was recorded historically. 458 T. C. G. RICH, B. EDWARDS & D. A. PEARMAN as ° Tete LeLy hye A a ane Se, TP-e gn LPR ae pS rin new Ka 2 AY re 4 ha eReE za pesat (Gilimzs, ie ele \ ed aaa ase Cite Se ree rosa rer a aie... A la i q) ed ( mS f es as conor ie aeons ye RRS Gas FIGURE 2. Hieracium portlandicum. A, Typical plant growing on a boulder. B, Habit of a larger plant growing in deeper soil. C, Lower side of rosette leaf. D, Upper side of rosette leaf. E, Upper side of stem leaf. F, Side view of capitulum. G, Involucral bract showing hairs types. H, Silhouettes of involucral bracts of H. portlandicum. 1, Silhouettes of involucral bracts of H. leyanum. Scale bars: A, 2 cm; B-F, 1 cm; G, | mm. Dele dt CrGaRich: HIERACIUM PORTLANDICUM 459 FIGURE 3. Distribution of Hieracium leyanum sensu stricto in South Wales and England. Records are for 1 km squares with symbols plotted at twice that size for clarity. @, 2003-2005. O, pre-1950. X, error. REFERENCES ALEXANDER, M. P. (1969). Differential staining of aborted and non-aborted pollen. Stain Technology 44: 117— 12D BARRETT, W. B. (1912). Contributions to a flora of Portland, with special reference to Limonium recurvum, C. E. Salmon. Proceedings of the Dorset Natural History and Antiquarian Field Club 33: 96-143. BOWEN, H. J. M. (2000). The flora of Dorset. Pisces Publications, Newbury. CLEAL, H. & RICH, T. C. G. (2004). An investigation of pollen viability in Hieracium. Unpublished report, National Museums & Galleries of Wales, Cardiff. HAINES, F. H. (1926). Report of the Natural History Sectional Committee of the Dorset Natural History and Archaeological Field Club for the year 1925. Dorset Natural History and Archaeological Field Club 47: 177-179. IUCN (2001). IUCN Red list categories and criteria. Version 3.1. (UCN, Gland. LEY, A. (1898). Two new forms of Hieracium. Journal of Botany 36: 6—7. - LEY, A. (1902). H. caledonicum, F. J. Hanb. Report of the Botanical Exchange Club of the British Isles 2:16. LEY, A., RIDDELSDELL, H. J. & LINTON, E. F. (1908). H. platyphyllum, Ley. Report of the Botanical and Exchange Club of the British Isles for 1907: 298. MILLS, J. N. & STACE, C. A. (1974). Chromosome numbers of British Plants, 2. Watsonia 10: 167-168. PUGSLEY, H. W. (1948). A prodromus of the British Hieracia. Journal of the Linnean Society (Botany) 54: \— 356. RIDDELSDELL, H. J., HEDLEY, G. W. & PRICE, W. R. (1948). Flora of Gloucestershire. T. Buncle & Co., Arbroath. SELL, P. D. & MURRELL, G. (2006). Flora of Great Britain and Ireland. Volume 4. Cambridge University Press, Cambridge. SELL, P. D. & WEST, C. (1968). Hieracium L., in PERRING, F. H. ed., Critical Supplement to the Atlas of the British flora. BSBI, London. WADE, A. E. (1970). The flora of Monmouthshire. National Museum of Wales, Cardiff. WIGGINTON, M. J., ed. (1999). British Red Data Books. 1. Vascular Plants. 3rd ed. Joint Nature Conservation Committee, Peterborough. ZAHN, K. H. (1921-1923). Compositae — Hieracium, in Engler, A. Das Pflanzenreich 77 (IV, 280): 1—1146. (Accepted April 2007) 460 T. C. G. RICH, B. EDWARDS & D. A. PEARMAN APPENDIX | - HISTORICAL AND RECENT RECORDS OF H,. PORTLANDICUM SP. NOV. FROM PORTLAND V.C. 9, DORSET Portland, 11 June 1910, W. B. Barrett (BM, Barrett (1912) cited it as ‘Durdle Pier to Churchope, plentiful. Sparingly in West Weare’) East Weare. Church Ope Cove, 2 September 1931, R. Meinertzhagen (BM); between Pennsylvania and the prison, June 1879, W. C. Medlycott (DOR); on the south side of the descent from the ... by Shepherds Dinner, 15 June 1886, J. C. Mansel-Pleydell (DOR); 20 June 1910, G. E. Fulleylove (CGE). The Verne Plateau, 1925, M. J. Andrews (Haines 1926; Bowen 2000). West Weare. 1885, W. B. Barrett et a/. (Bowen 2000); Chesil Beach, amongst quarry debris at the foot of the cliffs, near the neck, 20 July 1907, H. J. Riddelsdell (BM, CGE, NMW); 21 August 1911, C. E. Salmon (BM); Hallelujah Cove, 1925, M. J. Andrews (Haines 1926); limestone rubble, 4 June 1988, R. FitzGerald, S. L. Jury, J. Bevan, R. Pankhurst ef al. (B. S. B. I. field meeting; photograph in RNG as Fortuneswell, and incorrectly cited elsewhere as East Weare, pers. comm. R. FitzGerald 2005); 1989, H. J. M. Bowen (Bowen 2000); block boulder scree SY684730, 20 July 2003, B. Edwards & D. A. Pearman. APPENDIX 2 — RECORDS OF HIERACIUM LEYANUM SSS. V.C. 34, WEST GLOUCESTERSHIRE Six plants in roadside quarry between Birdlip and Crickley Hill, undated, H. J. Riddelsdell, and not refound again (Riddelsdell et a/. 1948). No voucher has been traced, but given that H. /eyanum is so distinctive and that Riddelsdell had collected it elsewhere, the record is accepted. V.C. 35, MONMOUTHSHIRE Tarren yr Esgob in “Breconshire and Monmouthshire” (Ley 1902; Wade 1970); still present in the Brecon part of this site but not seen recently at the extreme eastern end in Monmouthshire, if it ever indeed occurred in v.c. 35. The record plotted in ST59 in Sell & West (1968) is based on a specimen from rocks at the north point of Piercefield Park, 6 June 1900, A. Ley (CGE), determined by Sell & West, but which T. C. G. Rich believes to be H. pachyphylloides (Zahn) Roffey. Neither species was present on 5 July 2005, T. C. G. Rich. V.C. 36, HEREFORDSHIRE Red Daren, Hatteral Hills, 1892, 15 August 1893, 24 June 1896 and 9 June 1905, A. Ley (Ley 1898, CGE, NMW); cher | is also cultivated material from this site dated June 1896 anv June 1897 (BEL, CGE, MANCH, NMW). A specimen from Longtown, 31 August 1886, A. Ley (CGE) is assumed to be from this site. Not refound, 27 June 2005, T. C. G. Rich, or on the adjacent Black Daren. V.C. 42, BRECKNOCKSHIRE Craig Cerrig-gleisiad (as Craig Gledsiau) central upper part, | August 1890 and 9 July 1895, A. Ley (BM, CGE); 7 July 1896, F. J. Hanbury (BM); August 1951, C. West (?7MNE); Cliffs, 3 October 1962, B. Seddon (NMW); Cliff ledges SN961218-9, 4 July 1964, J. N. Mills (MANCH); Old Red Sandstone cliffs, SN961219, scarce, 24 June 2003, T. C. G. Rich & P. Hill (NMW). Craig Cwm-du (as Craig du), Senni Valley, 13 June 1893, A. Ley (BM, CGE); cliffs, SN943211, 22 July 2004, M. Porter (NMW). Craig y Cilau (as Craig Cille), undated, A. Ley (Ley 1898). This site has been extensively searched between 2000 and 2002 for Hieracium by T. C. G. Rich; presumed lost to the extensive quarrying. Cwm Tarrell, head of, 8 September 1888, and roadside rocks at head of, 9 and 10 July 1895, A. Ley (BM, CGE); 7 July 1896, F. J. Hanbury (BM); roadside, 13 June 1953, C. E. A. Andrews (BIRM); 22 June 1953, C. E. A. Andrews, J. R., & P. D. Sell (BIRM); July 1959, C. West (MNE?); damp cliff by road, SN9720, 18 June 1965, J. N. Mills & J. R. J. M. (MANCH); abundant on roadside and in quarry, SN973207, June 2004, T. C. G. Rich; this may be H. W. Pugsley’s site “roadside SW of Brecon’, 3 July +935 (NMW). HIERACIUM PORTLANDICUM 461 Darren Fach (as Cefn Coed, Dan-y-Graig, Vaynor and possibly Cefn Cil-Sanws), limestone cliffs, 7 June 1894, 8 June 1894 and 28 May 1896, A. Ley (BM, CGE, NMW); 17 June 1932, H. A. Hyde & A. E. Wade (NMW); 20 June 1932, A. J. Wilmott (BM); 10 June 1957, C. E. A. Andrews (BIRM); scattered along top of cliff, SO019107, 9 June 2000, T. C. G. Rich (NMW). Ley (1898) cites also ‘and other spots near Merthyr Tydfil’. Darren Lwyd (as Taren Llwyd), 24 June 1896, 15 June 1897 and 8 July 1898, A. Ley (BM, CGE). Two populations of 10 plants at SO243330 and c. 40 plants at SO241339, 27 June 2005, T. C. G. Rich (NMW). Dyffryn Crawnon, lime rocks at head of, 12 June 1893 and 11 July 1900, A. Ley (BIRM, BM, CGE); Sandstone cliffs, 27 May 1896, A. Ley & W. A. Shoolbred (NMW); 21 June 1953, C. E. A. Andrews (BIRM); July 1954, C. West (?CGE); 22 plants, old red sandstone crag, SO091154, 26 June 1975, P. D. Sell & L. Farrell (NCC rare species form). c. 45 plants, SO095149, 27 June 2005, T. C. G. Rich (NMW). Fan y Big, 31 July 1900, A. Ley (CGE). Not refound, T. C. G. Rich 2004. Sychbant, swallow hole near, 28 May 1896, A. Ley (NMW). About 300 plants on cliffs above and by cave, Cefn Sychbant, SN98570962, 6 July 2005, T. C. G. Rich (NMW). Tarren yr Esgob (also as Black Mountains), 12 July 1893, June 1896, 12 July 1897, 1 July 1898, 7 July 1898, 8 July 1898, 4 July 1901, 12 August 1903, 17 September 1907, A. Ley (BEL, BM, CGE, E, LIV, MANCH, NMW, some of which may be cultivated but is not necessarily marked as such); 29 July 1902, W. R. Linton (Set of British Hieracia no. 90; NMW, etc.); 9 June 1905 and 4 July 1907, A. Ley (BM, NMW); 23 July 1923, H. J. Riddelsdell (MANCH, NMW); 20 June 1933, C. E. A. Andrews (BIRM); Old Red Sandstone, June 1953, C. West (MNE”); 20 June 1953, J. E. Raven (E); 20 June 1953, J. E. Lousley (RNG); Scattered on top cliff, c. 30 plants at SO250308, 27 June 2005, T. C. G. Rich (NMW), but there are likely to be more extensive populations to the west on this large site. Watsonia 26: 463-468 (2007) 463 Conservation of Britain’s biodiversity: Hieracium cacuminum, Summit Hawkweed (Asteraceae) le sGaNiCn: Department of Biodiversity and Systematic Biology, National Museum of Wales, Cardiff CF 10 3NP and P. A. SMITH 128 Llancayo Street, Bargoed, Mid Glamorgan CF&1 8TP ABSTRACT Hieracium cacuminum, Summit Hawkweed, is a rare endemic Welsh hawkweed which has been confused with H. siluriense. A review of the records and field work showed that 240 plants are known in three sites; it was not refound in three other sites. It occurs on Old Red Sandstone cliffs and rocks in the Brecon Beacons. It is ‘Endangered’ under the IUCN Threat Criteria. KEYWORDS: Brecon Beacons, Endemic, Hieracium siluriense, [UCN Threat Criteria, Lectotype, Wales INTRODUCTION Hieracium cacuminum (Ley) Ley, Summit Hawkweed or in Welsh Heboglys y Copa, and H. siluriense (F. J. Hanb.) P. D. Sell, Silurian Hawkweed or in Welsh Heboglys y Silwriaid, are two Welsh endemic hawkweeds which have been confused historically as a result of mixed material being cited with the original des- cription of H. cacuminum by Ley (1895). Consequently, the distribution information in Ley (1895, 1909), Pugsley (1948) and Sell & West (1968) included both species. Sell & Murrell (2006) have recently clarified that the material on which H. cacuminum was based referred only to material collected in 1888 on the central Brecon Beacon cliff, and the other records cited belonged to H. siluriense. In order to clarify the distribution of H. cacuminum, a Red Data Book species (Wigginton 1999), and to provide an IUCN Threat Category as required under the Global Plant Conservation Strategy (Secretariat for the Conservation of *e-mail: tim.rich@museumwales.ac.uk Biodiversity 2002), the records and ecology of H. cacuminum were reviewed, along with fresh field surveys, and are reported in summary herein; full details are held by T. Rich and are available on request. Hieracium cacuminum was first described as H. diaphanum var. cacuminum by Ley (1895). A lectotype has been selected by P. D. Sell from material collected from a stream side, Brecon Beacons, 8 August 1888, A. Ley (herb. Ley, CGE) and is hereby designated by P.D.S. Hanbury (1904) used Ley’s epithet when he transferred the variety to H. vulgatum. Later Ley (in Linton 1905) transferred it to H. demissum, and then raised it to species status after the Scandinavian Hieracium expert G. A. H. Dahlstedt had seen material in 1907 (Ley 1909). Summary of synonymy: Hieracium cacuminum (Ley) Ley, J. Bot. 47: 51 (1909). angustatum (Lindeb.) Lindeb. subsp. cacuminum Zahn, in Engl., Pflanzenr. 76 (IV, 280): 471 (1921). =H. demissum Strémf. var. cacuminum Ley in W.R. Linton Brit. Hier. 74 (1905). =H. diaphanum Fries var. cacuminum Ley, J. Bot. 33: 86 (1895). =H. vulgatum Fries var. cacuminum Hanb. in Bab. Man. ed. 9, 258 (1904). Pugsley (1948) regarded H. cacuminum as an anomalous member of section 7ridentata. Sell & West (1968) and Sell & Murrell (2006) treat - it as a member of section Oreadea. The English and Welsh names are derived from the Latin cacumen = peak, extreme top. =F. 464 PACAGeRI@HES P. A. SMITH TABLE 1. REVISED HISTORICAL RECORDS OF HIERACIUM CACUMINUM V.C. 42 BRECON ‘Brecon Beacons’ (SO02) Brecon Beacons, central cliff 14 July 1886 Brecon Beacons, stream side 8 August 1888 Brecon Beacons 25 August 1890 Brecon Beacon, on the face of the cliff 26 August 1890 near the summit Brecknock Beacon (=Brecon Beacon), 26 August 1890 precipice of the Brecon Beacons, sandstone mountain 15 August 1894 cliff Brecon Beacons 28 July 1899 Brecon Beacons, rocks near the path 23 July 1902 between the two beacons (original locality) Cwm Tarrell (SN92) Cwm Tarrell, by a stream 24 July 1883 Cwm Tarrell August 1888 Cwm Tarrell 1890 Head of Cwm Tarrell, rocky 9 July 1895 mountainside Glyn Tarrell, cultivated June 1953 Glen Tarrell, rocky outcrop at head of — 10 July 1959 Taf Fechan (SOO1) Taf-fechan, sandstone cliffs 15 August 1894 Taf Fechan, | mile above the Ystrad 23 July 1900 Fan Gyhirych (SN8818) Y-fan-Gihirich 4 August 1898 Glyn Collwng (SO0519) Rocks at the head of Glyn Collwng 18 July 1908 (limestone)/ Collwng Daren, east side above railway V.C. 44 CARMARTHEN Llyn y Fan Fach (SN8021) Llyn y Fan Fechan, crags of 3 August 1899 Cultivated ex Brecon Beacons June 1898, 28 July 1899 A. Ley CGE JN SUES CGE (Lectotype), BM, NMW A Wey BM, Pugsley (1948) A. Ley BM A. Ley & W.H. CGE Purchas A. Ley & W.A. BM, CGE, LIV, NMW Shoolbred A. Ley BM, CGE, E, LIV, NMW; Set of British Hieracia no. 116 W. R. Linton BM, CGE, E, LIV, NMW; Set of British Hieracia no. 116; Linton (1903) AS Bey, BM Aeey, BM A. Ley BM, Hanbury (1894) Ae Ley. CGE, Ley (1909) PSDs Sell CGE C. West CGE W. A. Shoolbred NMW A. Ley CGE Aaey; CGE, Ley (1909) A. Ley CGE, Ley (1909) A. Ley CGE, Ley (1909) A. Ley BM, CGE, BEL, LIV IDENTIFICATION Hieracium cacuminum is illustrated in Figure 1. Key field key identification features separ- ating it from most other Hieracium species in South Wales are the flexuous stem with 3—6 stem leaves decreasing in size up the stem and without a clear basal rosette at flowering, and the few-headed inflorescence of capitula with dark involucres and very broad (1-7—-2:3 mm wide), obtuse outer involucral bracts. Vegetative plants have similar rosette leaves to the lower leaves on the stems of flowering plants, but where similar species (such as H. siluriense) were present and there was doubt about the identification, these were not recorded. HIERACIUM CACUMINUM 465 FIGURE |. Hieracium cacuminum and H. siluriense. A-K, Hieracium cacuminum. A, Whole shoot. B—G, Stem leaves from base showing decrease in size up stem (details of hairs not shown). H, First year rosette. I, Bud. J, Capitulum. K, Involucral bract. S, Involucral bract of H. siluriense. Scale bars 1 cm. Del. T. Rich. 466 TC) G RICH S& PTA] SME Hieracium siluriense differs from H. cacuminum in usually having at least a loose basal rosette with O—2(—3) stem leaves and narrower (to 1-7 mm wide), acute or subacute outer involucral bracts. The number of stellate hairs on the margins of the involucral bracts varies in both species, but in general there are more in H. siluriense. Full descriptions of both species are given by Sell & Murrell (2006). DISTRIBUTION Historical records were compiled from herbaria (BEL, BM, CGE, E, LIV, NMW), literature and the Hieracium database maintained by D. J eiMcCoshy} the revisedasreconds of ae cacuminum are given in Table |. Hieracium cacuminum has been recorded in six sites in two vice-counties in the Brecon Beacons in Wales, where it is endemic. The historical records were used to direct field work, principally carried out in July 2004. Voucher specimens have been placed in the Welsh National Herbarium (NMW). BRECON BEACONS (CENTRAL CLIFFS) 105 flowering H. cacuminum plants were found scattered along the cliff ledges on the NE facing side of Pen-y-fan on Old Red Sandstone between SO013215 and SO017212. They were often in clustered subpopulations, characteristically occurring on the sides of steep ‘alcoves’ formed by streams eroding the cliffs. The population estimate is a minimum as the crumbling sandstone cliffs are difficult to access safely. Hieracium siluriense is also present on the cliffs in greater numbers. Other crags of the central Brecon Beacons were also searched. Cribyn (SO0221) had some suitable ledges, with some H. siluriense and an unknown species. Craig Cwm Cynwyn (SO0220) had locally frequent H. siluriense and one clump of H. argenteum Fr. The rocks on the NE side of Fan y Big (SO0320) and Craig Cwarell (SO0420) were rather acidic and no Hieracium was seen. Corn Du (SO0021) had no suitable ledges. CWM TARRELL, CRAIG Y FRO Cliffs, rocks and gullies at the head of Cwm Tarrell were searched and H. cacuminum was found in two places on Craig y Fro. Nineteen flowering and c. 100 vegetative plants occurred on Old Red Sandstone cliffs by a waterfall at SN973203, and 4 flowering and 10 vegetative plants occurred on a small west-facing cliff at SN970206. At least five other Hieracium species were also present including H. siluriense. FAN GYHIRYCH The north-facing corrie has a low line of cliffs with a strongly calcifuge flora, and the only hawkweed seen was one patch of H. sparsi- folium Lindeb. GLYN COLLWNG Two flowering plants of H. cacuminum were found on the rocks on the west side of the waterfall of Nant Bwrefwr, immediately out- side the car park at SO055175, with H. cf. diaphanum agg. Two other Hieracium species were collected further upstream. No 4H. cacuminum was found elsewhere. TAF FECHAN The imprecise nature of the records made searching difficult, and no H. cacuminum was found in areas searched. Hieracium siluriense occurred at and below Neuadd Reservoir, and other species were found downstream in shaded woodlands. No H. cacuminum was seen on Graig Fan Ddu (SO0119) or Craig Gwaun Taf (SO0020). LLYN Y FAN FACH No H. cacuminum was found during extensive searches of the cliffs above the lake in 1999 and 2002, and it is probable that intensive sheep grazing has eliminated this and up to seven other hawkweeds which were recorded/ collected by A. Ley 100 years ago. The records are mapped in Figure 2. A total of c. 240 plants were seen in three sites in three hectads in one vice-county. It was not refound in three sites, but some of the localities are very imprecise and the original site may not have been found. BIOLOGY AND ECOLOGY Hieracium cacuminum is a polycarpic peren- nial which is dispersed by seed. Buds of two capitula were excised with scissors to remove the stigmas and anthers in a green-house, and covered; seeds were set in both excised and control buds, showing that reproduction is apomictic (voucher in NMW). The chromo- some number is not known, but it is likely to be polyploid. HIERACIUM CACUMINUM 467 J] © ‘ . Ox Se = 4 FIGURE -2. Distribution of Hieracium cacuminum. (@ 2004-2005. O pre-2004. X = errors, mostly referable now to H. siluriense). All populations of H. cacuminum occurred on low cliffs of Old Red Sandstone, either rooted in crevices or on immature soils derived from the sandstone. The ledges have vegetation 10-30 cm tall, with a moderate mixture of species. Typical species associated with H. cacuminum were Festuca ovina L. agg. (including F. vivipara (L.) Sm.), Solidago virgaurea L., Luzula sylvatica (Huds.) Gaudin, Deschampsia flexuosa (L.) Trin, Rhytidia- delphus loreus (Hedw.) Warnst. and Polytrichastrum alpinum (Hedw.) G.L. Sm. In a few places H. cacuminum is also associated with more calcicolous species such as Campanula rotundifolia L. and Thymus polytrichus A. Kern. ex Borbas. The upland cliffs of Pen-y-fan in which H. cacuminum occurs are probably best ascribed to the U16 Luzula sylvatica — Vaccinium myrtillus cliff ledge vegetation of the national vegetation classification (Rodwell 1992). All the sites are heavily grazed by sheep, and H. cacuminum is confined to ungrazed ledges and rocks. It is likely to be susceptible to grazing, like most other Hieracium species. Soil pH, measured with a pHep2 Hanna pocket-sized pH meter in a 50:50 mixture with distilled water of soil samples collected from around the roots, were pH 3-9 (ledge) and 7:1 (by waterfall) at Craig y Fro, and pH 3-7, 4:2 and 4:5 from ledges on Pen-y-fan. This range suggests that it is tolerant of both acid and calcareous soils. The soils were all damp but not waterlogged. The altitudinal range was c. 480 m at Craig y Fro to c. 650 m on Pen-y-fan. CONSERVATION Under the IUCN (2001) Threat Criteria, H. cacuminum qualifies as ‘Endangered’ (total population less than 250 individuals, popu- lation declining). The remaining populations on Pen-y-fan and Craig y Fro are probably toler- ably safe, as they are within the Brecon Beacons SSSI and SAC within the Brecon Beacons National Park. Although these designations offer no specific protection for H. cacuminum, the SAC is designated for the acidic and calcareous chasmophytic vegetation and the cliff ledge vegetation, in which H. cacuminum occurs. The two plants at Nant Bwrefwr are not in any designated sites, are close to a public car park and picnic area, and are highly threatened. The biggest threats to H. cacuminum are from rock-falls, which could eliminate local populations, and over-grazing. It is possible that the decline in the three sites is attributable to over-grazing, though without knowledge of the precise original localities this is difficult to assess. Forestry or development of reservoirs could also be responsible for the loss from Taf Fechan. A reduction in grazing throughout the Brecon Beacons would benefit this and other Hieracium species. Seed was collected from Craig y Fro and Pen-y-fan for the Millennium Seed Bank on 23 July 2004. No live collections are currently held in cultivation in botanic gardens. ACKNOWLEDGMENTS We would like to thank Joe Daggett and the National Trust, Andy Jones, David McCosh, Graham Motley, Mike Porter, Robin Walls and the Keepers of the Herbaria for access to libraries and specimens, and Peter Sell for allowing us to publish his lectotypification. The Countryside Council for Wales funded collation of the historical records. 468 Wa (Cy (Ge IeskClal ois IP, al, SIMU REFERENCES HANBURY, F. J. (1894). Notes on British Hieracia. Journal of Botany 32: 225-229. HANBURY, F. J. (1904). Hieracium, in Babington, C. C. (1904). Manual of British Botany. 9th ed. Gurney & Jackson, London. IUCN (2001). IUCN Red list categories and criteria. Version 3.1. YUCN, Gland. Ley, A. (1895). Short notes. Hieracium diaphanum Fr. var. cacuminum, n. var. Journal of Botany 33: 86-87. Ley, A. (1901). H. nitidum, Backh., var. siluriense, F. J. H. Report of the Botanical Exchange Club of the British Isles for 1900: 637. LEY, A. (1909). Brecon and West Yorkshire hawkweeds. Journal of Botany 47: 8-16 and 47-55. LINTON, W. R. (1903). Hieracium vulgatum, var. cacuminum. Report of the Botanical Exchange Club of the British Isles for 1902: 54. LINTON, W. R. (1905). An account of the British Hieracia. West, Newman & Co., London. PUGSLEY, H. W. (1948). A prodromus of the British Hieracia. Journal of the Linnean Society of London (Botany) 54: 1-356. RODWELL, J. S., ed. (1992). British plant communities. Volume 3. Grasslands and montane communities. Cambridge University Press, Cambridge. SECRETARIAT FOR THE CONSERVATION OF BIODIVERSITY (2002). Global strategy for plant conservation. Secretariat for the Conservation of Biodiversity, Montreal. SELL, P. D. & MURRELL, G. (2006). Flora of Great Britain and Ireland. Volume 4. Cambridge University Press, Cambridge. SELL, P. D. & WEST, C. (1968). Hieracium L., in, PERRING, F. H., ed., Critical Supplement to the Atlas of the British flora. BSBI, London. WIGGINTON, M. J., ed. (1999). British Red Data Books. I. Vascular Plants. 3rd ed. Joint Nature Conservation Committee, Peterborough. (Accepted January 2007) Watsonia 26: 469-472 (2007) 469 Rumex patientia L. x R. conglomeratus Murray, a newly described dock hybrid G. D. KITCHENER* Crown Villa, Otford Lane, Halstead, Sevenoaks, Kent TN14 7EA ABSTRACT A new hybrid, Rumex ~ philpii Kitchener hybr. nov. (Polygonaceae), between the neophyte R. patientia L. and native R. conglomeratus Murray, is described from a discovery in West Kent in 1978. KEYWORDs: Hybridisation, Patience Dock, Clustered Dock. INTRODUCTION On 9 July 1978, in the course of recording for the purposes of the Atlas of the Kent Flora (Philp 1982), a population of Rumex was encountered by E. G. Philp in a dried-up reed bed near Snodland, West Kent (v.c. 16), which included Rumex patientia and some evident hybrid plants. One of these specimens was identified as Rumex patientia x conglomeratus (the hybrid between Patience and Clustered Docks), and confirmed as such in 1989 by J. R. Akeroyd and D. H. Kent from material gath- ered on 21 July 1978 and held in MNE. Other material is Rumex patientia < obtusifolius, and includes one specimen which appears more sterile than is usual for that taxon. It has been speculated that this may be the product of complex hybridization in which a third species may also be represented, but there is insufficient evidence to conclude that it is other than R. patientia x obtusifolius. There appears to be no other record of R. patientia =< conglomeratus, whether in the British Isles or otherwise. The site of its discovery lies in the valley of the River Medway at TQ7161, where a dried-up reed bed approaches the river wall, the river being tidal at this point. The environs of the site have been subject to disturbance, through maintenance of drainage channels, gravel extraction and various industrial (and more recently, leisure) uses. In the 1970s, Rumex patientia formed a fairly isolated introduction here, with another colony near Grain, and with most other West *e-mail: geoffreykitchener!1@yahoo.com Kent records being located along the Thames Valley from the London boundary eastwards to Gravesend. It is possible that the frequency of this species is overstated in the records that were contributed to Philp (1982) (E. G. Philp, pers. comm. 2006), at the expense of the closely related Greek Dock, R. cristatus, which has since spread considerably. This, however, does not cast any doubt on the identity of the Snodland colony of R. patientia. The hybrid was distinctive by virtue of its high level of sterility. Even by late July, there was no evidence of seed formation and its tepals, while developing irregularly, did not include any which were fully formed, although the extent of development of some was such that it was possible to form an opinion in relation to the influence of putative parents. The tepals of R. patientia are entire, broadly ovate or sub-orbicular and may carry an inconspicuous tubercle. Those of Rk. conglom- eratus are also entire, but are lingulate and bear three conspicuous tubercles. The tepals of the hybrid were entire and varied from broadly ovate to lingulate, but tubercles were either small or mal-formed. The hybrid’s tepals were up to 3-5 mm long, intermediate between those of R. conglomeratus (2—3 mm) and R. patientia (S—8 mm). Local candidates for the parent other than Rk. patientia based on the possession of entire tepals would have been R. crispus, R. hydrolapathum or R. conglomeratus. Only the last-named of these is likely to have contri- buted the lingulate shape which is at one end of the spectrum of variation exhibited by the hybrid. R. crispus appears also to be ruled out as a parent by the insufficient crisping of leaf margins. In any event, observations of R. patientia x crispus elsewhere in West Kent in 2006 point to a more comprehensive maturing - of the flowering parts at least a month earlier as being normal. R. hydrolapathum offers a more plausible alternative as a parent, but would be expected to provide cuneate leaf bases, which 470 G. D. KITCHENER TABLE 1. CHARACTERS OF RUMEX PATIENTIA, R. CONGLOMERATUS AND THEIR HYBRID Rumex patientia < conglomeratus Rumex patientia Rumex conglomeratus Inflorescence Congested, upright branches; occasional leafy bracts towards most of their length base of inflorescence Tepals Entire Entire; 5—8 x 5-9(10) mm; 2-3 x 1-2 mm; broadly ovate to sub- _ lingulate; orbicular; generally bearing one small tubercle Ovate-lanceolate with subcuneate base Basal leaves Open, branches wide angled; leafy bracts present along bearing 3 large tubercles Oblong- to ovate-lanceolate with rounded base Somewhat loosely congested, with upper branches upright (Fig. 1A), lower branches more spreading; many narrow leafy bracts present at least in lower half of branches, sometimes extending into upper half (Fig. 1B) (Fig. 1D) Entire; generally imperfect but more developed tepals up to 3:5 x 2-5mm; broadly ovate to lingulate; occasionally with tubercles, somewhat elongate and mal-formed (Fig. 1C) Oblong- to ovate-lanceolate with subcuneate or rounded base were not present on this plant. The extent to which the hybrid demonstrated intermediacy between R. patientia and R. conglomeratus 1s set out in Table 1, but the key features are the tepal shape with its range of variation, the congested panicle inherited from R. patientia and the presence of leafy bracts subtending the flower whorls generally at least along the proximal half of the branches. DESCRIPTION Rumex * philpii Kitchener, hybr. nov. (Rumex patientia L. x R. conglomeratus Murray) Hybrida inter Rumicem patientiam L. et R. conglomeratum Murray, characteribus inter parentium characteres variantibus. Herba robusta erecta, altior quam 1 m, bracteas angustas foliaceas florum verticillos subtendentes saltem in ramorum florentium dimidio inferiore, nonnunquam ad ramorum extremitates, ferens; valvae fructiferae integrae, lingulatae vel late ovatae; semina omnino vel plerumque sterilia. Hybrid between Rumex patientia L. and R. conglomeratus Murray, with characters varying between those of the parents. Robust, erect herb, over | m high, bearing narrow leafy bracts subtending the flower whorls at least in the lower half of the flowering branches and sometimes to the ends of the branches; fruiting valves entire, lingulate to broadly ovate; seeds entirely or for the most part sterile. HOLOTYPUS: West Kent, v.c. 16, Snodland, in dried up part of reed bed, TQ7161, E. .G. Philp (MINE). There are three sheets marked Rumex patientia < conglomeratus in MNE, and that designated as holotype is sheet 2 of the gathering of 21 July 1978. The specific name is given in honour of Eric Philp, the finder of this hybrid, and who has done much to promote the understanding of the flora of Kent. DISCUSSION R. patientia is a native of south west Asia and south east Europe, and is widely but locally naturalised elsewhere in Europe (Jalas & Suominen 1979; Akeroyd, 1993) and in North America (Mosyakin 2005). Within the British Isles, it is scattered, but mainly present in the south east (Preston ef a/. 2002); although form- erly grown as a vegetable, it may also be a constituent of grass seed (Grob 1978). R. conglomeratus has a wide native presence across Europe and in west and central Asia, and is naturalised elsewhere in temperate areas. There is accordingly considerable overlap in distribution. RR. patientia 1s” mainly sruderalaere conglomeratus grows in damp, open habitats. Whilst there is potential for hybridisation where R. patientia grows on the banks of watercourses and so approaches the wetter habitats preferred by R. conglomeratus, their RUMEX = PHILPII 47] “0a % BPya ) FIGURE 1. Rumex x philpii. A. Inflorescence. B. Branch. C. Lower leaves. D. Fruits. E. Diagrammatic section through fruit. Rumex conglomeratus. F. fruit. Rumex patientia. G. Fruit. Scale bar for A-C = 1 cm. Scale bar for D and E= 1 mm. Scale bar for F and G= 1 mm. 472 flowering periods in the British Isles constitute an isolating mechanism, as they do not normally overlap (R. patientia, May to June; R. conglomeratus, July to September). Hence, although species within subgenus Rumex generally appear to hybridise freely, this is not a cross to be expected in normal circumstances. There is a parallel to be drawn in relation to the position regarding R. cristatus, which 1s sufficiently closely related to R. patientia for its separate specific status to have been queried by Stace (1997). Jauzein (1990) remarked on the failure of R. cristatus and R. conglomeratus to hybridise in a mixed group, where R. cristatus and R. crispus crossed readily. A similar disinclination to hybridise has been noted by the author in a large mixed colony of R. cristatus and R. conglomeratus at Rainham Marshes in South Essex, although this location has produced R. x akeroydii (Rumsey 1999) and hence demonstrated the feasibility of crosses between early and late flowering docks (in this case, R. cristatus and R. palustris). R. G. D. KITCHENER patientia 1s also recorded as having hybridised with R. palustris (R. x peisonis) in Austria (Rechinger 1933), which is further evidence that the isolating mechanism of normal flowering periods can be overcome. The extreme rarity of R. =< akeroydii and R. x peisonis, however, also suggests that overcoming such mismatches in flowering periods is a highly uncommon event. It appears unlikely that R. x philpii will have arisen through earlier flowering of R. conglomeratus, but rather that delayed or secondary flowering of R. patientia may have taken place, perhaps as a consequence of damage to the plant or disruption of the habitat. ACKNOWLEDGMENTS I am grateful to Eric Philp for arranging the loan of MNE material and for supplying information in relation to the find. Thanks are also due to Philip Oswald, for enhancing the Latin diagnosis. REFERENCES AKEROYD, J. R. (1993). Rumex L., in TUTIN, T. G. et al., eds. Flora Europaea 1: 99-107 (2nd ed.) Cambridge University Press, Cambridge. Gro, R. (1978). Identifikation um Rumex-Keimlingen. Seed Science Technology 6: 563-578. JALAS , J. & SUOMINEN, J. eds. (1979). Atlas Florae Europaea 4 Polygonaceae. Committee for the mapping of the flora of Europe and Societas Biologica Fennica Vanamo, Helsinki. JAUZEIN, P. (1990). Rumex cristatus DC. en France. Le Monde des Plantes 437: \-3. MOSYAKIN, S. L. (2005). Rumex, in Flora of North America, North of Mexico, 5 Magnoliophyta: Caryophyllidae, part 2. Oxford University Press, New York & Oxford. PHILP, E. G. (1982). Atlas of the Kent Flora. The Kent Field Club, West Malling. PRESTON, C. D., PEARMAN, D. A. & DINES, T. D. (2002). New Atlas of the British & Irish Flora. Oxford University Press, Oxford. RECHINGER, K. H. (1933). Vorarbeiten zu einer Monographie der Gattung Rumex II. Die Arten der Subsektion Patientiae. Repertorium specierum novarum regni vegetabilis 31: 225-283. RUMSEY, F .J. (1999). Rumex =< akeroydii — anew Dock hybrid. Watsonia 22: 413-416. STACE, C. A. (1997) New Flora of the British Isles (2nd ed.) Cambridge University Press, Cambridge. (Accepted January 2007) Watsonia 26: 473-476 (2007) 473 A reconsideration of Orobanche maritima Pugsley (Orobanchaceae) and related taxa in southern England and the Channel Islands Fee WINES E Department of Botany, Natural History Museum, Cromwell Road, London SW7 5BD ABSTRACT Sub-specific status is considered more appropriate for the plant currently known as O. minor var. maritima. Its distribution in south-east England is reviewed and revised. The distinctive yellow coastal taxon described from Jersey, originally as a form of O. ritro (= O. elatior), is recognised here as a variant of O. minor subsp. maritima and the necessary new combination made. KEYWORDS: British Isles. Orobanche, Taxonomy, Broomrape, INTRODUCTION Orobanche minor Sm. is throughout much of its British range a rather ephemeral species clearly often introduced in seed mixes and undeniably neophytic. Intra-populational vari- ation tends to be low and inter-populational differences great but without any geographical structure as one might expect of an inbreeding species which is routinely and randomly introduced. Populations in coastal situations in southern England have greater claims to native status and pose a more complex series of taxonomic and nomenclatural problems. Among the most distinctive of these plants, and that with the most coherent distribution and tightly defined host range, is the taxon described by Pugsley (1940) as O. maritima Pugsley. This is currently recognised as O. minor Sm. var. maritima (Pugsl.) Rumsey & Jury. Material ascribable to this taxon was first (incorrectly) recorded in the British Isles under the name O. amethystea Thuill. by the Rev. W. S. Hore (Hore 1845). The title of his paper indicates that he clearly had some doubts about its identity, not least because of apparent differ- ences in the lower lip of the corolla and the smaller sepals of his plant, but ultimately he seems to have become convinced that this was the correct determination. This view was clearly not shared by Borrer (see Pugsley 1940) but the plant was treated under this name as a *e-mail: F.Rumsey@nhm.ac.uk possible sub-species of O. minor by Syme (1866). He described its diagnostic features as a “corolla bent back into a quadrant in the lower third, the upper two thirds of the back nearly straight; [the] lower lip with the middle seg- ment conspicuously larger than the others.” Its host range and distribution were reported as “parasitical on Daucus in Whitesand Bay, Cornwall; near Plymouth, Devon; on the undercliff, S.E. of St. Margaret’s Bay , S. Kent; on Eryngium maritimum, near Cobo, Guernsey and St. Ouen’s Bay, Jersey.” One probable outcome though of this account was that Eryngium maritimum became fixed in the botanical public’s consciousness as a host to O. amethystea auct. Angl. non Thuill., or O. maritima as it was to become (Pugsley 1940) and maritime examples of the O. minor aggre- gate occurring on this host were uncritically accepted as being of this taxon. This confusion has been further perpetuated by the similarity in host to the true continental O. amethystea, a large reddish-brown plant closely allied to O. minor, primarily parasitic on Eryngium campestre, which has erroneously but under- standably been recorded as_ British subsequently, eg. Philp (1982). Pugsley (1940) neatly summarised much of the earlier confusion when recognising O. maritima aS a good species and he added several new distinguishing characters: the dark purplish colouration, an often bulbous stem- base, dense short pubescence, short broad- based bracts, shorter less acuminate sepals which were somewhat contiguous, 1.e. fused beneath the corolla, and a less deeply lobed stigma. He considered the host range of his new species to include Plantago coronopus and Ononis repens as well as Daucus, and first. made the linkage of this taxon with the coastal form of the wild carrot, “D. gummifer’ now regarded as D. carota subsp. gummifer (Syme) Hook. f., while deliberately excluding material from the Isle of Wight and Channel Islands parasitic on Eryngium maritimum. 474 THE TAXONOMIC STATUS OF OROBANCHE MARITIMA Of the additional characters cited by Pugsley (1940) when recognising this taxon at specific rank many are discontinuously variable and highly plastic; intensity of colouration is at least partially influenced by the nutrient status of the plant, itself controlled by the nature, 1.e. both the species and the vigour, of its host. Stem bulbosity is again dictated by the nature of the haustorial interface and this too 1s host- specific. Bract and sepal shape are highly and continuously variable and therefore not char- acters that can be used in isolation. Corolla shape (curvature of the back) is also difficult to quantify, highly variable, and fails to correlate with other supposedly diagnostic characters. The fusion of the stigma lobes is a condition which may be demonstrated by plants which show few or none of the other “maritima” characteristics and apparently may not always be shown by those that do (see Syme 1866, Plate MXVII). Likewise the degree of fusion of the sepals, a character that consistently defines such taxa as O. elatior and continental species such as O. clausonis Pomel, is variable within a single spike and the majority of the sepals examined on the holotype (Seacombe, Dorset, Pugsley 587 (BM) ) are un-fused. The sole remaining defining character is that of the shape of the lower lip of the corolla, with its pronounced rounded, less erose-edged, central lobe with large basal bosses; a feature difficult to see in herbarium material. Quantification and therefore identification of intermediates with the typical O. minor state is accordingly difficult. That said, on balance most material can be assigned to a taxon on the overall balance of the characters. This becomes increasingly difficult, however, at the southern and western end of O. maritima’s range in Cornwall, the Scillies and Channel Islands, where it apparently exhibits a broader host and habitat range, and at the disjunct eastern end of its range on the chalk cliffs and dunes of Kent. It is perhaps revealing that its primary host Daucus carota subsp. gummifer shows a similar if less marked eastward disjunction (Preston et al. 2002) and likewise intergrades morphologically with its more common inland counterpart in this area. It is accordingly tempting to suggest a closely linked co-evolutionary history for these taxa (Rumsey 1994). The areas in which we have the most difficulty discriminating taxa are those contact zones between var. maritima and other closely related plants. In the Purbeck limestone area of F. J. RUMSEY the Dorset coast from which the taxon was described by Pugsley, the plant is essentially isolated by habitat, but on lower coastal cliffs in proximity to dune grasslands and vegetated shingles to the west (Devon, Cornwall, the Channel Islands) there exists ever greater opportunity for gene flow from O. minor s. s. Confusion is also possible in south-east Kent with the extremely rare O. picridis F. W. Schultz, which occupies a similar ecological niche, if different host(s). It is clear from the error in Sitwell (1984) that this species is mistaken for O. minor var. maritima. Webb and Chater (1972) clearly regarded O. maritima as possessing some similarity to O. picridis: characters such as the straighter corolla back, more hirsute filaments, usually entire, not bidentate, calyx segments, etc. are common to both. On some well filled Victorian herbarium sheets from the St. Margaret’s area some individuals which otherwise appear to be O. picridis have shorter, more highly pigmented corollas, and shorter, if still filiform sepals (eg. St. Margaret’s, 1875, Bennett (BM)). A similar example, but more closely approaching O. minor s.l., was photographed by Philip Chantler near St. Margarets in June 2004. The narrow strip of coastal grassland abutting onto cultivated land east of St. Margaret’s also currently supports O. minor s. s. in arable margins and associated trackways and intermediate plants between this and O. picridis, and others which more closely approach var. maritima, are currently being investigated by C. Thorogood as part of his doctoral research. The pattern of distribution, degree of morph- ological separation and the potential gene-flow under sympatry, etc. demonstrated by this taxon, I now consider is best recognised by according it sub-specific level under O. minor. Orobanche minor Sm. subsp. maritima (Pugsley) Rumsey, stat. nov. Basionym: O. maritima Pugsley 1940 J. Bot., Lond. 78: 110. HOLOTYPE: Seacombe, Dorset. Pugsley 587, BM PARATYPES: Dover to Folkestone, 1882, Bennett (BM); Downderry, East Cornwall, 1876, Briggs (BM). [syn.: O. minor Rumsey & Jury]. var. maritima (Pugsley) O. minor subsp. maritima can best be distinguished from subsp. minor by the pronounced yellowish bosses on the lower _corolla lip, which has the middle lobe larger than the laterals and rather reniform in shape. OROBANCHE MINOR SUBSP. MARITIMA 475 The corolla is straight backed with a sharply inflected base, has short, often entire sepals and a broad-based rather triangular bract that does not exceed the flower. The plant is short, rarely exceeding 30 cm., the stem is puberulent, strongly pigmented and always bulbous at its base. DISTRIBUTION OF OROBANCHE MINOR SUBSP. MARITIMA IN KENT The distribution of O. minor subsp. maritima in Kent has been made uncertain due to confusion with other maritime forms of O. minor. The only reported chromosome count for QO. maritima (Hambler 1958) actually relates to the controversial O. minor population on the sand dunes of Sandwich Bay. Here, plant stature (height, corolla length) and pigmentation are highly variable, variation that appears in large part to be correlated with host identity. Plants with abnormally developed flowers, pedi- cellate, or with very reduced corollas and fused stigmatic surfaces, have also been seen here by the author. The records for this hectad (TR36) given by Stewart ef al. (1994) are therefore suspect. High chalk sea cliffs and the undercliff ledge/talus slope habitat do not extend northwards of TR34 and I have seen no unequivocal material that I would assign to subsp. maritima from TR35 or TR36. Hanbury and Marshall (1899) listed Syme’s record from St. Margaret’s (TR34); from the Undercliff at Lydden Spout, west of Dover (TR23), where found by Hanbury and a further record from the Dover area by Walton, that they thought perhaps referred to a different station. Hanbury and Marshall also note that “ Mr. Druce finds a specimen from Dover Cliffs in herb. Dillenius which he is disposed to place here”. In more recent times Philp (1982) only knew of a single small colony in tetrad TR23Z, presumably Hanbury’s Lydden locality. Since the year 2000, plants clearly referable to maritima have been seen on cliff ledges and cliff-tops in three separate hectads in E. Kent, v.c. 15, at the following locations: Lydden Spout (TR2939), Samphire Hoe (TR3039), Lighthouse Down, west of St. Margaret’s Bay (TR3644), and between Monument and St. Margaret’s Bay (TR3744). All populations are small and the total annual counts in recent years have not exceeded 60 plants and are often much lower. Cliff erosion in the St. Margaret’s area threatens the plants’ long-term survival immediately west of the Bay and the dense shading by trees of ledges immediately behind the bay is posing an additional threat. THE STATUS OF OROBANCHE MINOR VAR FLAVA The first recognition of what has come to be called O. minor var. flava E. Regel in British floras was made by Syme (1866). He noted after discussing “O. amethystea” “there are several other forms of O. minor which possibly deserve to be considered as sub-species; one occurs near Grand Havre, Guernsey [but see Druce’s (1911) comment], on Leontodon autumnale, which has the corolla curved like O. amethystea, but much shorter and denser spike, and the whole plant, including the flowers, is yellow”. As Syme correctly pointed out, the yellow-flowered taxon once present on coastal dunes in Jersey and Alderney (Pugsley 1940) is morphologically closer to subsp. maritima than to subsp. minor. Examples lacking anthocyanin are known in many Orobanche species and it has been suggested these may result from a single gene mutation (Rumsey & Jury 1991). These are best treated taxonomically as formae. The Channel Island plant, however, had subtle but distinct morphological differences, a different preferred host range (Asteraceae-tribe Lactuceae) and a discrete distribution. The rank of variety therefore seems appropriate but must be newly combined under subsp. maritima. The question then follows as to whether the name by which we have known this plant since Pugsley (1940) is correct. I consider that this taxon is not synonymous with O. minor Sm. var. flava Regel, or the later var. /utea Tourlet, both of which from their published descriptions (and geography) are almost certainly forms of O. minor subsp. minor which lack purple pigmentation. The earliest available epithet for the distinctive Channel Island plant would seem to be Beck’s “hypochoeridis”, a name with a troubled history. Material sent by Druce to Beck was apparently named by the latter as a form of O. ritro Gren. & Godr.; this name Druce then published without formal description, or citation of type, as forma hypochaeroides Beck (Druce 1907). Druce revised his opinion as to the rank of this taxon, recognising it as a variety (Druce 1911) but it still remained as a nomina nuda. This was appreciated by Beck (1922) who upon transfer of O. ritro into his O. major (= O. elatior Sutton) arguably validated 476 F. J. RUMSEY the name as O. major f. hypochoeridis, the authority for which was inaccurately cited by him as “(Druce) Beck”. The Channel Islands plant would thus become: Orobanche minor Sm. subsp. maritima (Pugsley) Rumsey var. hypochoeridis (Beck) Rumsey comb. et stat. nov. Basionym: O. major L. forma hypochoeridis Beck in Fedde, Rep. 18:36 (1922). HOLODVRE? WJerseya. st) Oueness Bay, aon Hypochaeris radicata L. and other composites, EAVe) ester-Garland Min herby Gai Dimice (OXF) [syn.: Orobanche minor Sm. var. flava angl. auct. non E. Regel]. The plant was apparently last recorded in Jersey in 1951(as O. ritro, A. G & F. W. Holder (Rumsey & Jury 1991) but the small population there, while possessing comparatively dense, round-topped inflorescences, differs in the other distinguishing features and furthermore is parasitic on legumes. Further work is needed to establish its identity and relationships. It is thus possible that Orobanche minor subsp. maritima var. hypochoeridis is now extinct. ACKNOWLEDGMENTS I am grateful to Chris Thorogood for his information on population sizes and locations in 2006, Dr. Helena Crouch for her useful comments on an earlier draft, Dr. Mark Spencer for assistance with nomenclature and access to the BM collections, and to the many other herbarium curators who have made their Orobanche specimens available to me over the last 20 years or more. s.n. (LIV!)). More recently it has only been recorded from Newport Docks (v.c. 35) REFERENCES BECK VON MANNAGETTA, G. (1922). Orobancheae novae. Fedde, Repertorium specierum novarum regni vegetabilis 18: 33-40. DRUCE, G. C. (1907). Notes on the flora of the Channel Islands. Journal of Botany, British and Foreign 45: 419-428. Druce, G. C. (1911). Short notes. Orobanche ritro Gren. & Godr. var. hypochaeroides. Journal of Botany, British and Foreign 49: 300-301. HAMBLER, D. J. (1958). Chromosome numbers in some members of the family Orobanchaceae. Botanical Journal of the Linnean Society 55: 772-777. HANBURY, F. J. & MARSHALL, E. S. (1899). Flora of Kent. F. J. Hanbury, London. Hore, W. S. (1845). Description of a species of Orobanche new to Great Britain, probably Orobanche amethystea, Thuillier. Phytologist 2: 239-240 PHILP, E. G. (1982). Atlas of the Kent Flora. Kent Field Club, Maidstone. PRESTON, C. D., PEARMAN, D. A. & DINES, T. D. (2002). New Atlas of the British and Irish Flora. Oxford University Press, Oxford. PUGSLEY, H. W. (1940). Notes on Orobanche L. Journal of Botany (London) 78: 105-116. RUMSEY, F. J. & JURY, S. L. (1991). An account of Orobanche L. in Britain and Ireland. Watsonia 18: 257— 295. RUMSEY, F. J. (1994). Orobanche minor var. maritima, in STEWART, A., PEARMAN, D. A. & PRESTON, C. D. Scarce Plants in Britain. J. N. C. C. Peterborough. SITWELL, N. (1984). Shell Guides to Britain’s Threatened Wildlife. London. STEWART, A., PEARMAN, D. A. & PRESTON, C. D. (1994). Scarce Plants in Britain. J. N. C. C., Peterborough. SYME, J. T. B. (1866) ed., English Botany, 3rd ed., vol. 6., Robert Hardwicke, London. WEBB, D. A. & CHATER, A. O (1972). Orobanche, in TUTIN, T. G., BURGES, N. A., HEYWOOD, V. H., Moore, D. M., VALENTINE, D. H., WALTERS, S. M. & WEBB, D. A. Flora Europaea, vol. 3. Cambridge University Press, Cambridge. (Accepted December 2006) Watsonia 26: 477-480 (2007) 477 An overlooked boreal clubmoss Lycopodium lagopus (Laest. ex Hartm.) Zinserl. ex Kusen. (Lycopodiaceae) in Britain FP. J RUMSEY= Dept. of Botany, Natural History Museum, Cromwell Road, London SW7 5BD ABSTRACT Lycopodium lagopus (Laest. ex Hartm.) Zinserl. ex Kusen. (syn. L. clavatum L. subsp. monostachyon (Grev. & Hooker) Selander), a widespread circum- arctic taxon, is reported from Britain for the first time. First collected in the 19th century, it had been dismissed as L. clavatum and its significance not recognised. The appropriate taxonomic rank for this plant has long proved controversial but where sympatric with L. clavatum it apparently maintains its distinctions, supporting the view that it deserves specific.rank. Its current status and British range are unclear but it is considered likely to be extremely localised and as an arctic disjunct element in our flora facing at least as great a threat as its declining lowland counterpart, not least through global warming and upland management practices. KEYWORDS: Lycopodium clavatum subsp. mono- stachyon, Cairngorms, Arctic Stag’s-horn Clubmoss, WACO OMVEC TONE INTRODUCTION During routine study of herbarium material at BM associated with the production of a revised checklist of the Pteridophyte flora of the British Isles and Ireland (Rumsey, in prep.), and the B.S.B.I. hybrids project, two specimens of what were recognised as the boreal form of Stag’s-horn Clubmoss (Lycopodium clavatum L. sensu lato) were detected. This plant was initially described as a variety of L. clavatum from Canada, under the name _ var. monostachyon, by Greville and Hooker (Hooker 1831) and independently from Scand- inavia as var. Jagopus by Laestadius (Hartman 1858); it was still treated at this rank by Rothmaler in the first edition of Flora Europaea (Tutin et al. 1964). More recently the consensus view has seen its elevation to sub- species, e.g. Jermy in Tutin ef al. (1993), Ollgard & Tind (1993), Kukkonen in Jonsell (2000), but frequently with indications that specific rank might be more appropriate. *e-mail: F.Rumsey@nhm.ac.uk However, amongst the recent major floras, only Wagner & Beitel (1993) in their account for the Flora of North America have accorded it specific status, a rank first published by Kuseneva (1953). In spite of the massive historical interest in Pteridophytes in the British Isles, this taxon would seem to have gone un-noticed here. The club-mosses were certainly less regarded than the ferns during the Victorian fern craze, perhaps as they were less amenable to culti- vation and were not treated in the majority of the fern works of the period. We thus have fewer descriptive accounts of minor morph- ological variants than with the true ferns, many of which have proved to be worthy of higher taxonomic rank, or have been shown to be of hybrid origin. Subsequently the failure to detect this taxon may, in part, be down to its controversial and lowly varietal status, in- accessibility of extra-British literature, and that common failing, our failure to look critically at apparently distinctive plants that require little effort to identify. In these respects it closely mirrors our treatment of another critical Boreal clubmoss Huperzia selago (L.) Bernh. ex Schrank & Mart. subsp. arctica (Grossh. ex Tolm.)A. & D. Léve (Corner eft al. 1998). DISTRIBUTION Lycopodium lagopus (Laest. ex Hartm.) Zinserl. ex Kusen. (Arctic Stags-horn Club- moss) has an Arctic circum-polar distribution and is also recorded in the Swiss and Austrian Alps and in the High Tatras in the Carpathians (Dostal 1984). In the sub-arctic it is a common plant of well-drained base-poor tundra slopes in nutrient-poor dwarf-shrub heath, sub-alpine birch forest and rather dry coniferous wood- land. In the Scandes it entirely replaces L. clavatum in the north, becoming less frequent as one proceeds south but extending to southern Norway (Aust-Agder) and as far south 478 as Dalarna province in central Sweden (Kukkonen 2000). It is frequent in Northern Finland south to Kainuu province and then extends at similar latitudes in a zone of fluct- uating width of 2—5° on the meridian through- out the Soviet arctic (Griffiths & Packer 1995), along the Bering Coast of Alaska into North America and Canada. It is also present in S.W. Greenland. In North America it extends as far south as southern Michigan (Wagner & Beitel 1993), again replacing L. clavatum in the north and with only a limited zone of sympatry, as also noted for the Soviet Arctic. In these contact zones may be found transitional plants with peduncles longer than their solitary strobilt. IDENTIFICATION The two taxa most noticeably differ when fruiting: L. clavatum, the widespread lowland plant has a long main fruiting axis, or peduncle which is >2:5 cm at maturity. It usually bears two, or more, strobili, each on short branches (pedicels) at the apex of the peduncle. In contrast, Lycopodium lagopus (Laest. ex Hartm.) Zinserl. ex Kusen. has strobili that are borne singly, and are sessile, or on a short peduncle (usually in British material adem EJ. Clement. 506 PLANT RECORDS tBrachyglottis ‘Sunshine’ (B. compacta * B. laxifolia) 135/66.comxlax. *62, N. E. Yorks.: cliff grassland, Robin Hood’s Bay, NZ952051, V. Jones, 2001. tRudbeckia hirta 135/78.1. *©60, W. Lancs.: tow path, Lancaster Canal, Lancaster, SD481668, J. H. Clarke, 2006, det. E. J. Clement. ©Helianthus annuus 135/79.1. *67, S. Northumb.: rubbish dump in layby off A69 near Throckley, NZ154677, R. & J. Bowyer, 2006. tHelianthus ~ laetiflorus (H. tuberosus < H. pauciflorus) 135/79.2xpau. *12, N. Hants.: in old chalk pit near roadside, East Woodhay, SU406601, R. P. Bowman, 1979, det. E. J. Clement, HCMS. tHelianthus rigidus Desf.135/79.rig. *12, N. Hants.: Greatham (SW), SU7730, A. B. M. Brewis, 1960. @ ©Bidens ferulifolia (Jacq.) DC. 135/81.fer. *62, N. E. Yorks.: pavement cracks, Higher Market Place, Kirbymoorside, SE695886, W. A. Thompson, 2005. *©63,S. W. Yorks.: Bolton Road, Bradford, SE169349, B. A. Tregale, 2003, det. E. J. Clement. @ tCoreopsis tinctoria Nutt. 135/82.tin. *©60, W. Lancs.: tow path, Lancaster Canal, Lancaster, SD481668, J. H. Clarke, 2006, det. E. J. Clement. @ tCoreopsis verticillata L. 135/82.ver. *12, N. Hants.: Church Crookham, near Gelvert Stream, SU831531, T. Dove, 1983, det. A. R. G. Mundell. A garden escape far from houses. It just survived the building of a new track but succumbed to birch scrub growth c.1988. @ ©Calotis hispidula (F. Muell.) F. Muell. 135/CAL his. *12, N. Hants.: wool adventive amongst young apple trees, Benham’s Fruit Farm, Blackmoor, SU7731, B. & J. Goater, 1969. @ tLiatris spicata (L.) Willd. 135/LIA.spi. *©63, S. W. Yorks.: Nailers Rough, Bradford, SE119359, B. A. Tregale, 2004, det. E. J. Clement. @ tMelampodium montanum Benth. 135/MEL.mon. *10, Wight: several self-seeded plants in full bloom in roadside pavement cracks, Shanklin, SZ5881, D. T. Biggs, 2006, det. E. J. Clement, photograph C. R. Pope. See BSBI News 94: 30 (2003). @ tOsteospermum jucundum 135/OST.juc. *11, S. Hants.: disturbed ground, Throop, SZ1095, R. M. Walls, 2006. tTagetes erecta 135/TAG.ere. *©39, Staffs.: dozens of plants on recently disturbed open ground near River Tame, Two Gates, SK211015, R. .D. Martin, 2006. tTagetes patula 135/TAG.pat. *©39, Staffs.: a few plants on disturbed ground, Wolverhampton, SO933997, P. Newton, 2000. tAponogeton distachyos 139/1.1. *12, N. Hants.: twenty plants established and probably increasing in River Blackwater on both sides of footbridge near Government Road, Aldershot, SU885519, C. R. Hall, 2004. Known here since 1997. tLysichiton americanus 147/2.1. *10, Wight: well established and spreading by streamside in wood, Saltmoor Copse, SZ5387, C. R. Pope, 2006. tCalla palustris 147/3.1. *12, N. Hants.: small plant in large flooded sand pit, MOD Longmoor, SU808305, A. R. G. Mundell, 2003, HCMS, herb. A.R.G.M. Specimen kept (herb. A.R.G.M.) Arum maculatum « A. italicum 147/5.1x2. | *12, N. Hants.: Ashford Lodge, Ashford Chace, SU746266, A. B. M. Brewis, 1964. Recorded as ‘hybrid’, listed under A. italicum subsp. italicum, with comment “Hybrid also seen, outside Ashford Lodge, at road junction [SU]746266’. ©Pistia stratiotes L. 147/PIS.str. *12, N. Hants.: Basingstoke Canal, Pondtail Flash and scattered in canal channel for 300m to E, in quantity, SU8253, C. R. Hall, 1995. GR inferred. It did not survive the winter. @ Juncus ambiguus 151/1.8. | *93, N. Aberdeen: flat floor of quarry once used for salt storage, Aikey Bray, W of Old Deer, NJ962478, S. D. S. Bosanquet & C. D. Preston, 2006. Juncus < kern-reichgeltii (J. effusus < J. conglomeratus) 151/1.26x27. *12, N. Hants.: The Warren, SE of Oakhanger, SU7735, F. Rose, 1989. *63, S. W. Yorks.: Maltby Low Common SK5491, G. T. D. Wilmore, 2006. tJuncus ensifolius Wikstr. 151/l.ens. *11, S. Hants.: pond, Fair Oak, SU483142, J. P. Poland, 2004, conf. E. J. Clement. *12, N. Hants.: Longstock Park Nursery, SU367388, B. Goater & A. R. G. Mundell, 2006, det. E. J. Clement, HCMS, herb. A.R.G.M. Growing as a weed in a pot of /ris pseudacorus.@ , Eleocharis palustris < uniglumis 151/3.1*3. *12, N. Hants.: River Test, Wherwell to Chilbolton, SU3840, F. Rose, 1984. 5 PLANT RECORDS 507 ©Cyperus eragrostis 152/11.2. *62, N. E. Yorks.: several plants at base of wall, Thirsk, SE434822, V. Jones & W. A. Thompson, 2006. tCarex vulpinoidea 152/16.6. *58, Cheshire: reclaimed sand quarry, Moore Reserve, SJ590860, G. M. Kay, 2004, det. A. O. Chater, herb. G.M.K. Carex divisa 152/16.13. *58, Cheshire: damp coastal field, Meols, SJ244908, E. F. Greenwood, 2006, LIV. Carex humilis 152/16.50. *10, Wight: several long-established patches in chalk grassland, Afton Down, SZ3685, P. D. Stanley, 2006, det. D. A. Pearman, herb. C. R. Pope. Previously overlooked! Carex x turfosa (C. nigra < C. elata) 152/16.67x68. *60, W. Lancs.: alder willow carr, with both parents nearby, Little Hawes Water, Yealand Redmayne, SD479768, E. F. Greenwood & M. S. Porter, 2006. See Wheldon & Wilson 1907. {Stipa neesiana 153/9.1. ©63,S. W. Yorks.: Bradford City Centre, SE164335, B. A. Tregale, 1995, det. E. J. Clement. Ist record since 1970. {Stipa gigantea Link 153/9.gig. *©63, S. W. Yorks.: Filey Street, Bradford, SE169327, B. A. Tregale, 2005, det. E. J. Clement. @ Ist record for B.I.? {Stipa tenuissima Trin.153/9.ten. *12, N. Hants.: several young plants seeded into gravel path, Longstock Nurseries, SU367389, B. Goater & A. R. G. Mundell, 2006, det. E. J. Clement, HCMS, herb. A.R.G.M. With abundant Polypogon viridis. © tOryzopsis miliacea var. thomasii 153/10.1. | *38, Warks.: 50 Brandon Parade, Leamington Spa, SP321659, J. W. Partridge, 2005, conf. species, det. variety T. B. Ryves, WARMS. Festuca < fleischeri (F. arundinacea < F. gigantea) 153/12.2x3 = *12, N. Hants.: in field at Coneycroft Bottom, Selborne, SU731335, P. D. Stanley, 1998, det. T. A. Cope, HCMS. Festuca arenaria 153/12.6. 10, Wight: sand dunes, Norton Spit, SZ3589, P. D. Stanley, 2004, det. M. Wilcox. Ist record since 1970. ~©Lolium multiflorum < L. rigidum 153/13.2xrig. *12, N. Hants.: Blackmoor area, SU73R, J. E. Lousley, 1960, det. E. E. Terrell. An article by A. Melderis states “When examining British material of Lolium at the British Museum (Natural History), Dr E. E. Terrell.... identified a specimen collected by J. E. Lousley at ....and distributed in his Wool Adventive Flora of Britain under the name L. rigidum, as L. multiflorum x rigidum, which has not been previously recorded from Britain’. @ +Vulpia myuros 153/14.3. | *68, N. Northumb.: at base of fence near police station, Wooler, NT992281, Q. J. Groom, 2006, herb. G. A. Swan. Puccinellia distans 153/16.2. *t23, Oxon: thirty plants on S side of A415 at Culham, SU505954, H. J. Killick, 2006. +Briza minor 153/17.2. *58. Cheshire: disturbed land, Heswall, SJ2680, E. F. Greenwood, 2006, LIV. {Briza maxima 153/17.3. 83, Midlothian: gutter and outer edge of pavement, Craigleith Hill, Edinburgh, NT228748, E. A. Pilling, 2006, conf. D. R. McKean, E, Ist record since 1970. {Poa palustris 153/18.10. *80, Roxburghs.: edge of pool, Banks of Tweed at Monksford Bank, NT583324, R. W. M. Corner, 2006, herb. R.W.M.C. Ist record in a natural habitat since 1913. Poa bulbosa 153/18.13. @®11, S. Hants.: gravel bank of car park, Hatchet Pond, East Boldre, SU369016, BSBI Field Meeting, 2005, det. M. W. Rand & R. M. Walls. Possibly introduced with the gravel used to make the barrier. Agrostis x murbeckii (A. capillaris x A. stolonifera) 153/39.1x4. *63, S. W. Yorks.: Valley sidings, Bradford, SE159345, B. A. Tregale, 2005, det. T. A. Cope. tPolypogon viridis 153/46.2. 38, Warks.: alley pavement, Clarendon Street, Leamington Spa, SP330662, J. W. Partridge, 2005, conf. T. B. Ryves, WARMS. Ist record since 1932. *58, Cheshire: pavement weed, Bowden, SJ759868, G. M. Kay, 2006, conf. A. Lockton & M. W. Rand, herb. G.M.K. *60, W. Lancs.: ditch, Westby landfill site, SD355311, P. Jepson, 2006, LIV. *Bromus secalinus 153/50.7. ®@©39, Staffs.: many on field margin, Alrewas Hayes, SK134145, M. E. Smith & D. Wall, 2005. ®@©39, Staffs.: scattered through large wheat field, Pound Green, SO757798, J. E. Hawksford, 2006. tAnisantha tectorum 153/52.4. ©10, Wight: sown lawn, St Mary’s Hospital, Newport, SZ4990, P. D. Stanley, 2004. 1st record since 1970. 508 PLANT RECORDS Elytrigia =< acuta nothosubsp. obtusiuscula (Elytrigia atherica < E. juncea subsp. boreoatlantica) 153/56.2x3a. 10, Wight: rather frequent in sand dunes, St Helen’s Duver, SZ6389, P. D. Stanley, 2006, det. T. A. Cope. Ist record since 1970. tCortaderia selloana 153/62.1. *38, Warks.: large clump in rough grass, Lea Marston purification lake, SP314950, M. A. Arnold, 2005, conf. J. W. Partridge. *62, N. E. Yorks.: one plant self-sown in pavement crack, Hutton Rudby, NZ467066, V. Jones & W. A. Thompson, 2006. tCortaderia richardii 153/62.ric. | *83, Midlothian: about 6 plants in roof guttering of house in Ferry Road, Edinburgh NT243759, D. R. McKean, 2005. tEchinochloa crus-galli 153/68.1. 65, N. W. Yorks.: landfill in silage field, West Burton, Bishopdale, SEO14877, C. Helm & D. Millward, 2006. Growing with other aliens. Ist record since 1970: *©81, Berwicks.: one hundred plants in potato field with soya bean waste manure, Hutton, NT909539, M. E. Braithwaite & W. D. Gill, 2006. tSetaria italica 153/70.ita. | *©81, Berwicks.: ten plants in potato field with soya bean waste manure, Hutton, NT909539, M. E. Braithwaite, 2006. tSetaria pumila 153/70.pum. *©81, Berwicks.: two plants in potato field with soya bean waste manure, Hutton, NT909539, L. W. Gaskell, 2006, det M. E. Braithwaite. tSetaria verticillata 153/70.ver. *©81, Berwicks:.: thirty plants in potato field with soya bean waste manure, Hutton, NT909539, M. E. Braithwaite, 2006. tDigitaria ischaemum 153/71.1. 12, N. Hants.: the dominant grass, partly buried in pure sand, beside numerous tracks used by military vehicles, Slab, Bordon, SU783355, A. R. G. Mundell, 2005, det. T. B. Ryves. Not extinct as stated in 1996 Flora of Hampshire. Really abundant, e.g. patch 15 m across at SU78323554, but on many other tracks in the area, mostly small non-flowering plants. . O©Digitaria sanguinalis 153/71.san. *62, N. E. Yorks.: several plants in pavement cracks, by church, Whip-Ma-Whop-Ma-Gate, York, SE605518, V. Jones & W. A. Thompson, 2006, herb. V. Jones. *67, S. Northumb.: Bedford Strut, North Shields, NZ355683, A. & G. Young, 2006, herb. G. A. Swan. ©Anemanthele lessoniana (Steud.) Veldkamp 153/ANE.les. *38, Warks.: four clumps in pavement, alley, Leamington Spa, SP318648, J. W. Partridge, 2005, det. T. B. Ryves, WARMS. Growing in nearby garden. © tPennisetum villosum R. Br. ex Fresen. 153/PEN.vil. *11, S. Hants.: road verge, Eastern Parade, Southsea, SZ6698, G. Hounsome, 2006, det. T. B. Ryves. @ Apparently Ist record for British Isles. Typha = glauca (T. latifolia x T. angustifolia) 155/1.1*2. *138, Warks.: established on eutrophic lakeside, Edgbaston Pool, Birmingham, SP054841, D. A. Broughton, 2006, conf. J. Southey. Natural England survey. *60, W. Lancs.: with both parents on margin of pond (eld gravel pit), N. Pond, Brockholes Quarry, Preston, SD579311, E. F. Greenwood, 2006. Pollen not stained with anthocyanin. Morphologically intermediate. tHemerocallis fulva 158/5.1. *12, N. Hants.: garden discard on wasteland, River Road, Yateley, SU808615, D. L. Brookman & A. R. G. Mundell, 2003. *93, N. Aberdeen: roadside, Old Meldrum, NJ833285, I. P. Green, 2006. Convallaria majalis 158/13.1. | £113(S), Channel Is. (Sark): in ditch outside garden wall, Le Petit Beauregard, WV458757, M. L. Long, 2006. Ist record for Sark. tChionodoxa forbesii 158/22.1. *12, N. Hants.: Hurstbourne Tarrant, Netherton Bottom, SU389549, P. Billinghurst, 1997. Presumably garden escape. tChionodoxa sardensis 158/22.2. *62. N. E. Yorks.: several self-sown plants on grassy bank, near church, Ingleby Greenhow, NZ582063, V. Jones, 1998. tChionodoxa luciliae 158/22.luc. *62, N. E. Yorks.: a few plants in woodland, Bassleton Wood, Thornaby, NZ446159, V. Jones, 2001. tMuscari armeniacum 158/23.2. *93, N. Aberdeen: woodand, Pitcaple, NJ7425, I. P. Green, 2006. tAllium roseum 158/24.3. *12,. N. Hants.: small patch (c. 12 plants) on roadside bank opposite entrance of Ashford Chace, SU740265, D. Ball, 1994. tAllium neopolitanum 158/24.4. *38, Warks.: clump 1 metre square in hedge, alley off Wathen Road, Leamington Spa, SP324667, J. W. Partridge, 2006, conf. J. R. Roberts, WARMS. tAllium subhirsutum 158/24.5. | *12, N. Hants.: Wishanger, SU838385, A. R. G. Mundell, 2003, det. B. Matthews, HCMS, herb. A.R.G.M. Site bulldozed a few weeks later, but some plants survived PLANT RECORDS 509 tAllium triquetrum 158/24.7. *38, Warks.: several clumps by footpath, Tachbrook Road, Leamington Spa, SP318649, J. W. Partridge, 2005, conf. E. J. Clement, WARMS. tAllium paradoxum 158/24.9. _*38, Warks.: more than 100 plants in abandoned garden, St Mary’s Road, Leamington Spa, SP326655, J. W. Partridge, 2005, conf. E. J. Clement, WARMS. tGalanthus nivalis < plicatus 158/32.1x3. *81, Berwicks.: one clump near River Tweed, Clinthill, NT605324, M. E. Braithwaite, 2006. tGalanthus elwesii 158/32.2. *39, Staffs.: grass verge, Wombourne, SO875930, C. B. Westall, 2006. Present for many years on the roadside, sheltered by road signs from mowing. tGalanthus woronowii 158/32.wor. *12, N. Hants.: single very large clump beside boundary path on west side of Tadley Common, SU606621, |. & W. G. Helyar & A. R. G. Mundel, 2003, det. R. Baker, HCMS. Shown to A. R. G. Mundell by finder W. G. Helyar. Determined in situ in flower on 21 Feb 2004 by invited Galanthus expert. tNarcissus poeticus 158/33.3. *67, S. Northumb.: Howford Banks, NY9166, A. & G. Young, 2006. Has been present for at least 10 years. tNarcissus cyclamineus 158/33.6. *11, S. Hants.: woodland, Trodds Copse, Ampfield, SU420227, P. Marston, 2006. In NE finger of woodland, having spread from a nearby garden. tErythronium dens-canis 158/ERY.den. *67, S. Northumb.: Jesmond Dene, Newcastle, NZ26NE, A. & G. Young, 2004. tSisyrinchium californicum 159/2.3. *©39, Staffs.: three plants in paving cracks on Hundred Acre Road, NE of Pheasey, SP077968, M. W. Poulton, P. Reade & I. C. Trueman, 2006. tSisyrinchium striatum 159/2.5. *12, N. Hants.: Sheet, SU7524, A. Knapp & D. Nelson, 1999. tIris germanica 159/5.1. *67, S. Northumb.: Seaton Sluice, NZ337768, J. L. Durkin, 2005. tIris orientalis 159/5.7._ *10, Wight: well-established in roadside ditch, Brook, SZ3883, M. J. Burnhill, 2006, det. C. R. Pope. Iris foetidissima 159/5.8. *160, W. Lancs.: roadside verge, Silverdale, SD473759, P. Jepson, 2006. *t£81, Berwicks.: well-established clump in policies, The Hirsel, NT830409, M. E. & P. F. Braithwaite, 2006. 83, Midlothian: railway embankment, 200 m east of Bonnington Bridge above walkway, Edinburgh, NT264763, H. S. McHaffie & D. R. McKean, 2005.1st record since 1970. tCrocus speciosus 159/8.8. *©Ol11, S. Hants.: made earth bank near Brockishill Road, Bartley, SU298117, M. W. Rand, 2005. Introduced with landfill to remake bank of Forest car park. *12, N. Hants.: N side of track on grassy bank outside fenced garden, Pamber Heath, SU610618, W. G. Helyar, 2001, conf. A. R. G. Mundell. *62, N. E. Yorks.: a few plants in rough grassland, Grey Towers, Nunthorpe, NZ537133, V. Jones, 2001. tFreesia < hybrida 159/12.1. *©63, S. W. Yorks.: waste ground, Thorpe Salvin, SK5082, W. A. Ely, 2004, conf. J. R. Comley. Epipactis phyllanthes var. vectensis 162/3.7. 10, Wight: ancient woodland on chalk, Eaglehead Copse, Ashey, SZ5887, P. Jupp, 2005, det. A. J. Richards. First appeared in 2003 but confirmed 2005. Ist record since 1970. xGymnaglossum jacksonii (Gymnadenia conopsea < Coeloglossum viride) 162/16<17.1*1 *12, N. Hants.: Noar Hill, Selborne, SU7431, M. Oates, 1975. One plant in late 1970s noted in management plan appendix. Dactylorhiza < hallii (D. maculata x D. praetermissa) 162/18.2x4. *12, N. Hants.: Conford Moor, West Fen, SU8133, F. Rose, 1992. Dactylorhiza incarnata subsp. coccinea 162/18.3b. *63, S. W. Yorks.: Woodhall Quarry, near Calverley, SE197353, B. A. Tregale & G. T. D. Wilmore, 2002. Dactylorhiza < wintoni (D. incarnata < D. praetermissa) 162/18.3x4. Ne lantse: Humbly Grove area, N of Powntley Copse, SU74C, J. E. Lousley, 1980. Recorded as ‘Dactylorchis incarnata x praetermissa = D. < wintonii’. Dactylorhiza ~< latirella (D. incarnata < D. purpurella) 162/18.3~5. 64, Mid-W. Yorks.: with both parents in flushed marshy area beside outflow stream from Malham Tarn, altitude 385 m, Malham, SD893658, P. P. Abbott, 2006, conf. N. Barrett. Ist record since 1970. Dactylorhiza purpurella = D. traunsteinerioides 162/18.5x7. | *62, N. E. Yorks.: one plant in calcareous marsh, Sand Dale, Dalby Forest, SE858848, V. Jones, 2003, conf. R. D. Bateman. @ 510 PLANT RECORDS ~ Orchis purpurea 162/20.5. *12, N. Hants.: a fine plant with three flowering spikes and a dead one from previous year in Isle of Wight woods, Porton Down, SU251373, L. Delve et al., 2003, conf. A. R. G. Mundell. Ophrys apifera 162/23.3. *75, Ayrs.: on re-vegetated bit bing, Barony Colliery, New Cumnock, NS525218, B. Laney, 2003, conf. D. A. Lang. Ist confirmed record. See BSBJ News No. 95: 30 (2004). Still present in 2005. Photograph with vice-county recorder. Watsonia 26: 511—526 (2007) 511 Obituaries VERA GORDON 1918-2006 With the death of Vera Gordon on 14 Sept- initially rather awesome, with its goodly ember 2006 the Liverpool botanical community and the North West of England more widely have lost someone who for very many years has been a pivotal figure, just as the B.S.B.I. too has lost one of its longest-serving vice- county Recorders as well as one of its newest Honorary Members. Born on 25 May 1918, Vera was brought up in Liverpool’s northern suburbs, first in Bootle and later in Waterloo, and continued to live at the same address in the second of those for almost her entire adult life, To the warmth, helpfulness and lively humour for which that city is renowned she added the specially personal qualities of immense physical energy and a tirelessly enquiring mind. Both her parents and her only brother were keen ramblers and through that she was introduced to the country- side at an early age, acquiring an interest in wild flowers during family holidays in the Lake District and North Wales. The South Lancashire sand dunes, with their magnificently rich flora, also lay within easy walking distance of the family home. From primary school in Bootle she won a scholarship to the grammar school there, and she would surely have gone on to university had she been born half a century later. As it was, her formal education ended at sixteen and she went to work in the accounts department of a local firm of dry-cleaners. When the Second World War broke out she fondly eyed the Women’s Land Army as her way of contrib- uting to the war effort but she was deeply disappointed that her accounting experience proved to have earmarked her inescapably for similar office work in the Royal Ordnance Department. The obverse of that coin, however, was that she acquired the security of the Civil Service thereby, which subsequently enabled her to transfer to a career more to her liking as administrative officer in the local magistrates court, where she remained till her retirement. Meanwhile, her father’s membership of a vhotographic society had brought him the acquaintance of Dr C. Theodore Green, the author in 1933 of a second edition of the Flora of the Liverpool District, and at the latter’s instance Vera was encouraged to join the city’s long-standing Botanical Society. Barely eighteen then, she must have found that body sprinkling of Edwardian veterans who had built up over the years a breadth of expertise quite normal in their day, but by then becoming all too rare. The doyen, W. G. Travis, a collab- orator in his youth of the by that time legendary J. A. Wheldon, was typical in being as well- versed in bryophytes, lichens and microfungi as he was in flowering plants and ferns; for thirty years he had been compiling on the Society’s collective behalf a much-needed flora of South Lancashire, the densely-populated (and long well-botanised) vice-county 59. A young enthusiast with energy to spare was never- theless more than welcome, and the very next year she found herself elected to the Society’s Council and the year after that one of two Honorary Secretaries. Sixty-two years later she was still serving in that second capacity, a record of continuity interrupted only by an eventual term as President in 1952-1953 followed by three years as Vice-President. Throughout that exceptionally long period she played the principal part in holding that small society together, for without her efficiency and friendly informality it might well have petered out. In 1950, on the top of a swaying double- decker bus bound for the advertised venue of one of the Society’s field meetings, for which we were the only two to have braved the heavy rain, Vera learned of the recrudescence that the Botanical Exchange Club had recently under- gone in its new guise as the BSB.I. In common with many field botanists in the North, the Liverpool ones had long held aloof from the BEC on account of its lingering identification in their eyes with the 19th century tradition of reckless collecting; once Vera joined the B.S.B.I. however, and began passing around her copies of the new BSBI publications, all the other leading members speedily followed her example. Thereafter Vera was a frequent attendee, and always active participant, at BSBI field meetings, enlivening them with her infectious laughter and exemplary persistence in recording however atrocious the weather. One that proved partic- ularly memorable for her was that in West Cork in 1964. To get there, she had to make a lengthy journey by bus on which to her delight she found herself engulfed in inimitable Irish rustic chatter; then, once the meeting was under 2 OBITUARIES way, her sharp eyes spotted an unfamiliar plant with small white flowers which was to turn out to be another native Sandwort, Minuartia recurva, new to the British Isles list. By then her BSBI involvement had been substantially extended by her appointment as Recorder for South Lancashire, in which increasingly onerous capacity — coinciding as it did with the 1950s Distribution Maps Scheme — she was to serve for thirty-six years. Additionally, when a_ short-lived tier of ‘District Secretaries’ was introduced in 1960 to deal with the growing flood of enquiries of a less strictly scientific nature, she willingly shouldered responsibility for that secondary task in the group of vice-counties that made up the ‘Mersey’ Province. Around the same time the teaching of plant taxonomy at Liverpool University experienced a marked revival, consequent upon the appointment to the staff of Dr Vernon Heywood and his very active involvement in the massive Flora Europaea project. This intensified the need for a guide to the local flora for placing in the hands of students, and as a result the long-accumulating Flora of South Lancashire, which had been languishing because of the latterly frail health and then decease of Travis, had new life vigorously breathed into it. Vera was the natural choice as secretary of a new committee charged with the daunting task of using the manuscript left by Travis as the basis for a volume of a more modern character that was more realistically publishable. To her also fell much of the time- consuming checking of the many post-1939 records and the rounding-up of additional ones that were found to be needed, though that formed only one ingredient in the eventually seven years of combined effort out of which the much-modified Flora at last emerged into print, in 1963, with Vera’s name justly featuring alongside those of Heywood and Dr J. P. Savidge as joint editors. Such was the ability and care Vera had displayed in that prolonged undertaking that the University saw her as the obvious person to revitalise and curate its herbarium; but on being sounded out she reluctantly concluded that, with a non-transferable Civil Service pension in prospect, such a move involved too great a sacrifice financially. The University was not to forget her, however, for two decades later, in 1987, it conferred on her an honorary M.Sc. Legend has it that the University’s imposing Daimler sent to her home to convey . her to the degree ceremony created almost more of a sensation locally that the honour itself. Further recognition followed, In 2001 Vera finally allowed the Liverpool Botanical Society to elect her to honorary membership, and four years after that the BSBI honoured her similarly — appropriately at an A.G.M. held in Wales, over the north of which she had spent a great part of her life enquiringly striding. It was as a great walker, indeed, that Vera will probably be remembered by the greatest number of people above anything else. As the years went by. Britain proved too confining for her energies and many parts of the world, as distant as Australia and as remote as the Caucasus, attracted her as well. Often that was with a botanical group, in which her untiring assistance and increasingly very wide knowledge was ever at her companions’ disposal; but botanizing, with its unavoidably slow pace, was not enough to absorb her seemingly inexhaustible energeticness and she varied those occasions with more arduous challenges such as the famously testing Appalachian Trail. Other favourite outlets were Scottish dancing (at which she was accomplished enough to become a teacher of it at evening classes), gardening (alpines were a speciality) and photography, this last an enthusiasm passed on to her by her father and at which, like him, she came to excel. Alert and reasonably active almost to the end, Vera was able to attend an indoor meeting of her beloved *L.B.S.’ as late as six months before her death. Four further months were then spared her before finally having to exchange her house for a nursing home near Ormskirk, where, not long afterwards, her death took place. Her extensive herbarium of British plants, together with numerous speci- mens collected from around the world, had been donated by her to Liverpool Museum (now World Museum Liverpool) (LIV) some years before. Her large collection of 35 mm slides has joined those, at the wish of her executors, who have also presented to the library of the Liverpool Botanical Society most of her botanical books. For many of the details in this account I am indebted to Vera’s brother, Ron Gordon, and LBS members, in particular Susan Taylor, Joan Vincent and Eric Greenwood. DAVID ALLEN OBITUARIES ROBERT ALAN FINCH 1939-2006 Dr Bob Finch, who died of cancer on 18 December 2006, was by profession a plant cytologist and by inclination a botanist whose wide-ranging interests included fungi, lichens and bryophytes as well as vascular plants. His particular speciality was in the genus Leontodon, of which he contributed accounts to the standard reference works. Born on 20 April 1939, Bob showed an interest in natural history from a very early age. This interest was encouraged by one of the masters at Hastings Grammar School, Douggie Brightmore, a keen botanist. Bob’s records of flowering plants appeared annually in the records section of the Hastings and East Sussex Naturalist from 1957 until 1975 (excepting only 1972—1973). It was in this journal that he published his first paper at the age of only 19, a survey of the vascular plants and bryophytes of six roadside clinker heaps, undertaken in 1956— 1957 (Finch 1958). Clinker was used in those days for gritting roads in winter. Bob went on to Pembroke College, Oxford, where E. F. Warburg, then Britain’s leading bryologist as well as a one of the authors of ‘CTW’, was another major influence. In 1963, soon after completing his degree, Bob published a de- tailed paper on the bryophytes of the Hastings area (Finch 1963). Bob had arranged to follow his degree with a research studentship under Warburg’s Supervision, but the head of department, C. D. Darlington, insisted that Bob should be his student, threatening to make departmental life unpleasant for both him and Warburg unless he did so. Bob felt that he had to comply with Darlington’s wishes, although it would be difficult to imagine a less congenial supervisor for the mild-mannered and unassuming Bob. Indeed Darlington told Bob outright that he was stupid — it was only when he read a biography of Darlington 40 years later that Bob discovered, to his great relief, that Darlington had written in his diary that “At the age of 18 most of the world seemed stupid and annoyed me; at 24 I know it is stupid and it ceases to worry me” (Harman 2004). Despite the temperamental gulf between them, Bob completed a successful D.Phil. thesis at Oxford on the cytology of the genus Leontodon (Finch 1966). He later wrote it up as a paper during a year’s temporary lectureship at Newcastle (Finch 1967). During the fieldwork for his thesis he rediscovered the hybrid between L. hispidus and L. saxatilis, a plant hitherto reported in Britain only by another cytologist, K. B. Blackburn. Bob left Newcastle for a job as a plant cytogeneticist at the Plant Breeding Institute, Trumpington, near Cam- bridge, in 1967. He was recruited as a member of a new team set up to study barley, and during the course of his career he was co-author of over 30 papers on the cytology and genetics of barley. He continued to work in the Cytogenetics Department until he was made redundant in 1985, one of 42 departures in a year in which the PBI was being prepared for privatisation. At Cambridge, Bob was able to pursue his interest in Leontodon with Peter Sell. Together, they undertook the difficult job of preparing an account for Flora Europaea, producing a radical revision in which many existing species were reduced to subspecies (Finch & Sell 1976a, 1976b). Bob had contributed the account of Leontodon to Stace’s Hybridization and the Flora of the British Isles (Finch 1975) andielatenene sandmsketcr sScllardescrbedie hispidus x saxatilis as L. x vegetus in Sell & Murrell’s (2006) Flora of Great Britain and Ireland. Bob also made use of the large living collection of Hieracium which had been built upmby, Peter SellMand yal Westin the Cambridge Botanic Garden to investigate the cytology of apomixis in the genus. His observations are summarised in Sell & Murrell’s account of Hieracium. Bob was a very keen bryologist who collected two species new to Britain, Tortula freibergii in the Hastings area and Bryum valparaisense on St Agnes, Isles of Scilly (Preston 2007). Bob and his wife Jessica usually took their annual holiday on St Agnes, where he also. rediscovered the lichen Pseudocyphellaria aurata at a time when it was feared extinct in Britain. He and Jessica shared other interests, including wildlife sound- recording and the poetry of the Northampton- shire poet John Clare. Bob was an old-fashioned figure, quite out of sympathy with the prevailing attitudes of the late twentieth and early twenty-first centuries. He was modest and unassuming, incapable of pushing himself forward. At bryological meetings he was prepared when necessary to 514 devote all his time to beginners, helping them with a cogent exposition of the characters of the mosses they encountered. He would, however, never venture an opinion without outlining the reasons why he considered that he was really unqualified to express a view. Nevertheless, he could not conceal his wide- ranging botanical knowledge and a shrewd scientific intelligence. He was very much the absent-minded academic. The methodical, met- iculous and cautious approach to scientific problems which served him well as a botanist was also applied to the lesser problems of everyday life. There can be few who, like Bob, feel the need to draw a sketch map of the position of his car in the carpark before entering a motorway service station. (He was, rather surprisingly until one remembered his OBITUARIES perfectionism, a committed member of the Institute of Advanced Motorists.) Thus, every- day tasks tended to be time-consuming and he was not always the most punctual of men. These foibles greatly endeared him to the friends who met him regularly on the field meetings of the British Bryological Society’s Cambridgeshire and Norfolk & Suffolk groups, and provided the basis of many stories. Had his talents been combined with a more driven personality he would surely have achieved even more than he did, but it was clear from the large attendance at his funeral that it was just this character which had endeared him to his friends, and he introduced many younger people to the natural world he loved. I have never met anyone quite like Bob; he was a unique character and is already greatly missed. REFERENCES FINCH, R. A. (1958). A list of the flowering plants and mosses found on roadside clinker heaps. Hastings and East Sussex Naturalist 9: 1-3. FINCH, R. A. (1963). The bryophytes of the Hastings area. Hastings and East Sussex Naturalist 9: 185-191. FINCH, R. A. (1966). Chromosomal and morphological variation in the British flora. D.Phil. thesis, University of Oxford. FINCH, R. A. (1967). Natural chromosome variation in Leontodon. Heredity 22: 359-386. FINCH, R. A. (1975). Leontodon L. In: STACE, C. A., ed. Hybridization and the flora of the British Isles, pp. 429-430. London: Academic Press. FINCH, R. A. & SELL, P. D. (1976a). Leontodon L., in HEyYwoop, V. H. (ed.), Flora Europaea notulae systematicae ad Floram Europaeam spectantes no. 19. Botanical Journal of the Linnean Society 71: 241— 248. FINCH, R. A. & SELL, P. D . (1976b). Leontodon L., in TUTIN, T. G.-et al. (eds), Flora Europaea Volume 4 Plantaginaceae to Compositae (and Rubiaceae), pp. 310-315. Cambridge: Cambridge University Press. HARMAN, O. S. (2004). The man who invented the chromosome: a life of Cyril Darlington. Cambridge, Massachusetts: Harvard University Press. PRESTON, C. D. (2007). Obituary. Robert Alan Finch B.A., D.Phil. (1939-2006). Journal of Bryology 29 (in press). SELL, P. & MURRELL, G. (2006). Flora of Great Britain and Ireland. Volume 4 Campanulaceae—Asteraceae. Cambridge: Cambridge University Press. C. D. PRESTON OBITUARIES 515 FRANCIS ROSE 1921—2006 Francis Rose died on 15 July 2006, aged 84. He was unquestionably one of the finest field botanists of his generation and his knowledge of the flora of western Europe had few rivals. Francis was born in south London in 1921 and he frequently told how his interest in botany was first awakened by a naturalist grandfather who took him for country walks at the early age of six. He claimed to possess a photo of himself, aged eight, collecting plants in the Alkham Valley in Kent, clutching a specimen of what was apparently Mentha rotundifolia. Although there was no biology taught at his Roman Catholic school Saint Francis Xavier College, he went up to London University to read Botany in 1938, first at the Chelsea Polytechnic and then Queen Mary College, from where he graduated in 1941. In 1947, after wartime work testing explosives at Woolwich Arsenal and a stint of teaching Engineering Science at Gravesend Technical School he arrived at Sir John Cass College as lecturer in botany. Two years later he moved to Bedford College, where he stayed for the next fifteen years. Here he developed his early research interest in the ecology of lowland bogs and fens; he was awarded his PhD in 1953 for his thesis on Researches on the Floral Ecology of British Lowland Bogs and Heaths (Rose 1953). At this time he and his wife, Pauline Wendy, whom he had married in 1943, were living at East Malling, from where he explored the Kent and Sussex countryside, frequently in the company of the late E. C. (Ted) Wallace and R. A. (Ron) Boniface. He had already embarked upon a project to produce a new flora of the county, as none had appeared since the publication of Hanbury & Marshall’s Flora of Kent in 1899 (Rose 1949, 1955). His publication of the Biological Flora of Orchis purpurea, 1n which he noted morphological differences between populations on either side of the Stour valley, had been an early contribution to the Kentish Flora (Rose 1948). An account of the remnant fens around Sandwich followed in 1950 (Rose 1950) and at the same time he had embarked upon a three part bryophyte Flora of the county, which he completed in 1951 (Rose 124), ISSO, IOSD Francis maintained a continuing interest in mire ecology and was particularly fascinated by the East Anglian spring fens with their high number of what he regarded as late-glacial relic bryophyte species. With the assistance of his research student, David Bellamy, he carried out comprehensive surveys of Redgrave and Lopham Fens, which paved the way for their declaration as National Nature Reserves (Bellamy and Rose 1961). During this period he was closely involved with the emerging importance of the Field Study Council field centres. He co-directed, with Francis Butler, the University of London’s Certificate of Proficiency in Field Biology and had particularly close relations with Juniper Hall and Flatford Mill Field Centres. Orchids remained a particular interest. His very first paper described a new British species of Epipactis, E. vectensis, later submerged within E. phyllanthes (Brooke & Rose 1940), and he formally described the northern form of the Fragrant Orchid as a separate subspecies, Gymnadenia conopsea ssp. borealis (Rose QOL). In 1964 Francis moved to King’s College London as senior lecturer in Biogeography. He was appointed University Reader in 1975, a post that he occupied until his retirement in 1981. His botanical interests were essentially those of plant geography and phytosociology. He would not have claimed to be a classical taxonomist nor was he an experimental ecologist. He was fascinated with the interpretation and meanings of distribution patterns and this was combined with a formidable competence as a field botanist and an encyclopaedic knowledge of the west European flora. Many would argue that his most influential contribution to British botany is The Wild Flower Key, first published in 1981 and continuously in print for over twenty-five years until the 2nd edition appeared shortly before his death, revised and updated by Clare O’Reilly. Easily the most popular and successful of plant field guides, it broke new ground with the incorporation of vegetative keys arranged by habitat. The success of these was yet further evidence of his extraordinary familiarity with the plants in the field. This was followed in 1989 by what had originally been intended as a companion volume, Colour Identification Guide to the Grasses, Sedges, Rushes and Ferns of the British Isles and north-western Europe. \t was unfortunate that the relatively expensive, large format, hard-back edition put it beyond the reach of many students, as it, too, is still the best colour guide to these groups, 516 OBITUARIES including as it does Claire Dalby’s technically brilliant plates of the non-Carex Cyperaceae. Francis also became involved with that iconic series, the Observer’s Books, revising British ferns (1965), British grasses, sedges and rushes (1965), British wild flowers (1978) and, with Kenneth Alvin, Lichens (1977). It was natural that he would be drawn into the interest in disjunct distributions that had been stimulated] bya ythe BsS-Blks) sothe conference report, particularly in relation to south-east England (Rose 1957). He developed the idea that distribution patterns were often explicable in the context of post-glacial history. Certain areas of the chalk had higher con- centrations of local species than others and he noted that these were often in the vicinity of what could have acted as ‘refugia’ habitats. habitats that would have remained open during the Atlantic forest maximum. Similarly, the well-known concentration of oceanic species in the wooded ravines of the Central Weald, such as Hymenophyllum tunbrigense and Dryopteris aemula, were relics of the Atlantic period, surviving in a cool damp microclimate similar to that of the western seaboard. He explored all the known outcrops of the Wealden sand rocks, refinding most of the bryophyte species recorded by W. E. Nicholson at the beginning of the last century and discovering new colonies of Hymenophyllum. He also demonstrated the widespread occur- ence of Dryopteris aemula in these deep ravines, a plant that Wolley-Dod described as rare (Rose 1952; Pentecost and Rose 1985; Richardson, Rose & Rich 1985; Rose 1997). Francis was also intrigued by the odd disjunct distributions shown by some species of the chalk of northern France and that of Britain that were absent from the North and South Downs but reappeared again north of the London Basin. His comparisons of the flora of S. E. England and northern France appeared in a number of papers in the Bulletin de la Société de Botanique du Nord de la France during the 1960’s with his friend Prof. Jean-Marie Géhu. With Géhu’s encouragement he also was intrepid enough to produce a paper on the identification of north-west European Salicornia! (Rose 1989) Francis moved to Liss in the early 1960s, which was a convenient distance from the new field station at Rogate that King’s College had recently opened. It was also from about the mid-1960s that his increasing interest in lichens developed, encouraged by the late T. D. V. (Dougal) Swinscow and Peter James. This coincided with a further involvement in the use of plants as bio-indicators of past and present environmental conditions. With David Hawks- worth, he was one of the first to demonstrate the value of lichens growing on tree trunks as sensitive indicators of specific levels of atmos- pheric sulphur pollution. (Rose & Hawksworth 1970; Hawksworth, Rose & Coppins 1973; Hawksworth and Rose 1976). He also showed that forest lichens were not only sensitive to air quality. Certain species were only to be found in ancient undisturbed woodlands and this enabled him to construct a series of ‘indices of ecological continuity’, a technique that he later extended to higher plants. This work had important consequences for our interpretation of the nature of what Oliver Rackham has termed the ‘wildwood’, the presumed pre-agricultural forest cover of Britain. The richest sites were those with ancient trees set amid pasture grassland, such as old deer parks and parts of the New Forest, rather than continuous closed woodland. In this he anticipated by more than twenty-five years the recent theories of Franz Vera on the role of large herbivores. His higher plant lists of ancient woodland indicator species are now widely adopted in woodland conservation assessment. Originally produced as a series of regional lists for use in lowland England, each list consisting of a hundred species, they inevitably prompted a debate on the objectivity of such indices as well as less successful attempts to produce similar indices for other habitats, such as old grassland (Hornby & Rose 1987; Rose 1999). The move to Hampshire meant that the New Forest, as well as West Sussex were now much more accessible. There is no question that the New Forest held a special place in Francis’ affections. He regarded it as the finest remaining example of lowland deciduous forest in western Europe. He was a strong supporter of the moves to establish the National Park and was hugely influential behind the scenes in helping to ensure sensitive management of the woodlands. Francis was the leading authority in western Europe on the lichen epiphytes of old forest and his 1974 paper, with Peter James, on the corticolous and lignicolous species of the New Forest was a major contribution. The move to Hampshire also resulted in the plans to produce a Flora of Kent moving down the priority list and in the event it was never written, although he retained strong links with OBITUARIES SIF i the county. However, work in Hampshire resulted in The Flora of Hampshire, with Lady Anne Brewis and Peter Bowman, published by Harley Books in 1996. Five introductory chapters contain a masterly account of the vegetational history and ecology of the county, in the same way that his account of the ecology of Sussex in his Atlas of Sussex mosses, liverworts and lichens with Rod Stern, Howard Matcham and Brian Coppins (1991) is by far the best that there is. The latter was published as a separate by the Booth Museum in 1996. In all, Francis published over 140 books and papers, but arguably his most influential con- tribution lies in the countless unpublished reports for the Nature Conservancy and its successors, for local authorities and the Wildlife Trusts. The files of the old English Nature contain dozens of his reports that resulted in the original notification of a substantial proportion of all the Sites of Special Scientific Interest in southern England. The raw material for all of this output resides in the evidence of the prodigious amount of time that he spent in the field and recorded in his field notebooks and record cards. The 200 or so notebooks, commencing in 1944 and containing about 250,000 individual records, Francis Rose photographing Military Orchid in the Chilterns in June 2004 (Photo: D. Streeter). are held by the National Museum of Wales, together with the bulk of his herbarium, totalling over 18,000 specimens, and photographic slide collection. The note-books have been transcribed by Judith Church and computerised by the Sussex Biodiversity Records Centre. The rest of his archive (mostly lichens) is held by the Natural History Museum, and that relating to Kent by the Maidstone Museum. In 1955 Francis was closely involved in the establishment of the Kent Field Club. For a number of years he had been _ botanical secretary of the South-Eastern Union of Scientific Societies, an archaic survivor of the heyday of Victorian natural history. As Director of Field Studies of the KFC, he established a number of long-term surveys, notably into the population dynamics of the Lady Orchid, Orchis purpurea, on the North Downs above Maidstone. He remained a staunch supporter of local natural history and was President of the Somerset Natural History and Archaeological Society in 1987. Francis was deeply committed to the cause of conservation. He was a keynote speaker at the seminal conference held at the London Zoo in 1958 which established the County 518 OBITUARIES Naturalists’ Trusts in south-east England. He was the first Honorary Secretary of the Kent Trust and its Chairman from 1959-1965. He was an honorary member of both the Sussex and the Kent Trusts and a council member of the Hampshire and Isle of Wight Trust. He was a founder member of Plant Life and served on its Council. He campaigned vigorously for the proposed South Downs National Park until shortly before his death. Francis joined the B.S.B.I. in 1944 and served on both the Council and_ the Conservation Committee. He also served on the Council of both the British Bryological Society and the British Lichen Society, being elected President of the latter for 1980-1982. He was elected an honorary member of all three societies, the only British botanist ever to be so honoured. He was appointed M.B.E. in 2000 and received the Wildlife Trust’s Christopher Cadbury medal in the following year. In 2003 the Royal Botanic Gardens, Kew established the Francis Rose Reserve at Wakehurst Place, the first nature reserve in Europe to be dedicated to bryophytes, ferns and lichens. His name is celebrated in two species of lichens; Phyllopsora rosei and Porina rosei. It is perhaps easy to forget that for the whole of his professional life, Francis was first and foremost a university academic. But his students won't forget, as a letter in the current edition of Plantlife testifies (Spring 2007). He was an inspirational teacher and had an extraordinary gift of making even the common-place exciting and every fresh encounter with a familiar plant seem like a first meeting. His knowledge was encyclopaedic, not only about the plants themselves but about all that bore on their existence; art, history, culture, folklore. And he loved talking about them as anyone familiar with his telephone conversations will recall! He was a larger than life character, great company with a sometimes wicked sense of humour as the story of the famous telegram sent to the late J. E. Lousley by Francis and Richard Fitter on their discovery in 1956 on the discovery of his secret Chilterns site of the Military Orchid, Orchis militaris, bears witness. Although, unless prompted, lunchtime might well be overlooked, opening time rarely was! Throughout his life he enjoyed the support of his wife, Pauline Wendy. In his message to his 80th birthday conference at Cardiff he wrote that, “without her help and encouragement over sixty years I would not have achieved very much’. To her and his four children we extend our condolences. I would like to thank David Harper and Tim Rich for reading the draft of this obituary. DAVID STREETER Publications of Francis Rose (1921-2006) Prepared by T. C. G. Rich | 1940 Brooke, B. J. & Rose, F. (1940). A new British species of Epipactis. Journal of Botany 78: 91-89. 1942 Rose, F. (1942). A note on the rediscovery of Orchis simia in Oxfordshire. Journal of Botany 80: 102-104. 1945 Rose, F. (1945). Wild orchids of Britain. Discovery 6(7): 207-211. 1946 Rose, F. (1946). The vegetation of the Weald. South eastern Naturalist and Antiquarian 51: 1-6. 1948 Rose, F. (1948). Orchis purpurea Huds. Biological Flora of the British Isles. Journal of Ecology 36: 366-377. 1949 Rose, F. (1949). A bryophyte flora of Kent. I. Introduction and ecological notes. Transactions British Bryological Society 1: 202-210. Rose, F. (1949). Botanical records for metropolitan Kent 1942-1949. London Naturalist 1949: 15— 20. Rose, F. (1949). The flora of Kent.-Address to the Botanical Section, S.E.U.S.S. South-eastern Naturalist and Antiquarian 54:39-40. Rose, F. (1949). Excursions, Society of the British Isles: 40-45. 1947. September 5-8. SETH (Kent)., Year Book Botanical BIBLIOGRAPHY 519 1950 Rose, F. (1950). A bryophyte flora of Kent. II. Hepaticae. Transactions British Bryological Society 1: 225-265. Rose, F. (1950). The East Kent fens. Journal of Ecology 38: 292-302. 1951 Rose, F. (1951). A bryophyte flora of Kent. HI. Musci. Transactions British Bryological Society 1: 427-464. Rose, F. (1951). A note on Atrichum undulatum and A. angustatum. Transactions British Bryological Society 1: 481-484. Rose, F. (1951). Distribution maps of Kent plants. (Exhibit). In: The Study of the Distribution of British Plants. BSBI Conference Report no. 2: 76-79. BSBI, London. 1952 Rose, F. (1952). Annual reports. Botanical Section. South-eastern Naturalist and Antiquarian 55: XVIlI-XXIl. Rose, F. (1952). “Atlantic” species in the flora of the Weald. South-eastern Naturalist and Antiquarian 57: 18-23. Rose, F. (1952). Annual reports. Botanical Section. South-eastern Naturalist and Antiquarian 57: XVII-XIX. Rose, F. (1952). A bryophyte flora of Kent. 1V. Addenda. Transactions British Bryological Society Di Dae 1953 Rose, F. (1953). A survey of the ecology of the British lowland bogs. Proceedings of the Linnean Society 164: 186-211. Rose, F. (1953). Botanical records in Kent, Surrey and Sussex 1952-1953. South-eastern Naturalist and Antiquarian 58: 18—24. 1954 Rose, F. (1954). Botanical records for Kent, Sussex and Surrey 1953-1954. South-eastern Naturalist and Antiquarian 59: 2\—25. (DSS Rose, F. (1955). The New Flora of Kent. Presidential address to the Botanical Section. South- eastern Naturalist and Antiquarian 60: 9-19. 1956 Rose, F. (1956). Field Meetings, 1955. May 27th to 30th, 1955. Dover. Botanical Society of the British Isles Proc. 2: 176-178 Rose, F. (1956). Trientalis europaea L. Botanical Society of the British Isles Proc. 2:21. SST Rose, F. (1957). Vegetation-history and environmental factors in the London area. London Naturalist 37: 29-40. Rose, F. (1957). The importance of the study of disjunct distributions to progress in understanding the British flora. Progress in the study of the British flora. BSBI Conference report, no. 5: 61— 78 1958 Rose, F. (1958). Exhibition Meeting, 1957. Dryopteris aemula in S.E. England. Botanical Society of the British Isles Proc. 3: 100-101. 1959 Rose, F. (1959). Sussex records for 1958. S.-E. Naturalist and Antiquarian 63: 14—16. 1960 Rose, F. (1960). The rediscovery of Orchis simia Lam. in Kent, 1. Foreword. Transactions Kent Field Club 1: 50. Rose, F. (1960). Botanical records for Kent, 1955—S8.Vascular plants. Transactions Kent Field Club 1: 56-65. Rose, F. & Laflin, T. (1960). Kent bryophytes. Transactions Kent Field Club 1: 66-78. Rose, F. (1960). Records for 1960. Bulletin Kent Field Club 6: 8-11. Rose, F. (1960). The vegetation and flora of the Weald. Botanical Society of the British Isles Proc. 4: 96-97. Rose, F. (1960). Tilia platyphyllos Scop. in Kent. Bulletin Kent Field Club 5: 29. Rose, F. & Gehu, J. M. (1960). Comparison floristique entre les comtés anglais du Kent et du Sussex et le département fran¢gais du Pas-de-Calais. Bulletin de la Société de Botanique du Nord de la France 13: 125-139. 520 BIBLIOGRAPHY Rose, F. & Gehu, J. M. (1960). L’excursion de la ‘Botanical Society of the British Isles’ dans le Nord de la France; son apport a la connaissance de la flora et de la vegetation du Pas-de- Calais. Bulletin de la Société de Botanique du Nord de la France 13: \—12. 1961 Rose, F. (1961). The Burham Down Survey: |. Vegetation in 1956. Transactions Kent Field Club 1: 119-125. Bellamy, D. J. & Rose, F. (1961). The Waveney-Ouse valley fens of the Suffolk-Norfolk border. Trans. Suffolk Naturalists’ Soc. 11: 367-385. 1962 Rose, F. (1962). Reports of Recorders: Botanical. Bulletin Kent Field Club 7: 9. 1964 Rose, F. (1964). Compte rendu des Herborisations. II. Le Sud-Est de l’ Angleterre (Comtés de Kent et Sussex). Bulletin de la Société Botanique de France 111, 90e Session: 30-37. Rose, F. & Gehu, J. M. (1964). Essai de phytogéographie comparée. La végétation du Sud-Est de P Angleterre et ses analogies avec celle du Nord de la France. Bulletin de la Société Botanique de France 111, 90e Session: 38—70. Rose, F. (1964). Contribution pour une flore de Bryophytes de Nord de la France. Bulletin de la Société Botanique de France 111, 90e Session: 209-238. 1965 Rose, F. (1965). Comparison phytogeographique entre les pelouses craueuses du Meso- — Xerobromium des vallies de la Basse-Seine, de la Somme, de |’Authie, de la Cauche, de la Caesta Boulonnaise et du Sud-Est de |’Aangleterre. Revue des Societeies Savantes de Haute- Normandie (Sciences) 37: 105-109. Rose, F. (1965). The Observer’s book of British ferns. Revised edition. Frederick Warne & Co, London. Rose, F. (1965, 1974). The Observer’s book of British grasses, sedges and rushes. Revised edition. Frederick Warne & Co, London. 1966 Rose, F. (1966). Reports of field meetings held in 1965. 2nd October — Oldbury Hill. Bulletin Kent Field Club 11: 29-31. Lichens by F. H. Brightman. 1967 Rose, F. (1967). Reports of field meetings held in 1966. 23rd April — Biddenden area. Bulletin Kent Field Club 12: 1-111. Rose, F. (1967). Reports of field meetings held in 1966. 23rd October — Hythe area. Bulletin Kent Field Club 12: i1i-v. Rose, F. (1967). Eriophorum gracile Roth new to Ireland. /rish Naturalists’ Journal 15: 361-362. 1968 Rose, F. (1968). Reports of field meetings held in 1967. 14th October — Fairlight, East Sussex. Bulletin Kent Field Club 13: 35-38. Rose, F. (1968). Woodland ecology and conservation. Journal of the Arboricultural Society (publication untraced). 1969 Barkman, J. J., Rose, F. & Westhoff, V. (1969). The effects of air pollution on non-vascular plants. In: Air pollution. Proceedings of the First European Congress on the Influence of Air pollution on Plants and Animals. Wageningen 1968. Centre for Agricultural Publishing and Documentation, Wageningen, pp. 237-241. Rose, F. & Bailey, R. H. (1969). Lichens from the Forest of Dean — new or otherwise interesting records. Journal Gloucestershire Naturalists Field Club 20: 77-79. Rose, F. & Hawksworth, D. L. (1969). A note on the lichens and bryophytes of the Wyre Forest. Proceedings Birmingham Natural History and Philosophical Society 21: 191—197. Rose, F. (1969). Reports of field meetings. 20th April — Combwell Wood, Goudhurst. Bulletin Kent Field Club 14: 15-18. 1970 Rose, F., Hawksworth, D. L. & Coppins, B. J. (1970): A lichenological excursion through the north of England. Naturalist [Hull] 95: 49-55. Rose, F. (1970). Lichens as pollution indicators. Your Environment 1: 185-189. Rose, F. (1970). Plant life section (c. 60,000 words) In: Shell Natural history of Britain, ed. M. Burton. M. Joseph, London. BIBLIOGRAPHY 521 Rose, F. & Hawksworth, D. L. (1970). Qualitative scale for estimating sulphur dioxide air pollution in England and Wales using epiphytic lichens. Nature 227: 145-148. McClintock, D. & Rose, F. (1970). Cornish heath in Ireland. /rish Naturalists’ Journal 16: 387— 590) 1972 Rose, F. (1972). Floristic connections between South-east England and North France. Taxonomy Phytogeography and Evolution. BSBI Conference Report no.12: pp. 363-379. nSWs Rose, F. (1973): Detailed mapping in south-east England. In: B. W. Ferry, M. S. Baddeley & D. L. Hawksworth (eds.): Air Pollution and Lichens. University of Toronto Press, Toronto, pp. 77— 88.. Hawksworth, D. L., Rose, F. & Coppins, B. J. (1973): Changes in the lichen flora of England and Wales attributable to pollution of the air by sulphur dioxide. In: B. W. Ferry, M. S. Baddeley & D. L. Hawksworth (eds.): Air Pollution and Lichens. University of Toronto Press, Toronto, pp. 330-367. Rose, F. (1973). A mobile lichen — in which grid square? Bulletin British Lichen Society 33: 13. 1974 Rose, F. & Wallace, E. C. (1974). Changes in the bryophyte flora of Britain. In: Hawksworth, D. i ed., The changing flora and fauna of Britain. pp. 27-46. Systematics Association Special Volume no. 6. Academic Press, London. Hawksworth, D. L., Coppins, B. i, & Rose, F. (1974). Changes in the British lichen flora. In: Hawksworth, De ed., The changing flora and fauna of Britain. pp. 47-78. Systematics Association Special Volume no. 6. Academic press, London. Rose, F. & James, P. W. (1974): Regional studies on the British lichen flora I. The corticolous and lignicolous species of the New Forest, Hampshire. Lichenologist 6: 1—72. Rose, F. (1974). The epiphytes of oak. In: The British Oak. Its History and Natural History. Ed. M. G. Morris, & F. H. Perring. BSBI Conference Report no.14: 250-273. E. W. Classey, Faringdon. IO75 Rose, F. (1975). The lichens of the Gwaun Valley. Nature in Wales 14: 186-189. Rose, F. (1975). The vegetation and flora of Tycanol Wood. Nature in Wales 14: 178-185. 1976 Rose, F. (1976). Lichenological indicators of age and environmental continuity in woodlands. In: Brown, D. H., Hawksworth, D. L. & Bailey, R. H. Lichenology: progress and problems. Pp 279-308. Systematics Association Special volume no. 8. Academic press, London. Hawksworth, D. L. & Rose, F. (1976). Lichens as pollution monitors. Institute of Biology series no. 66. Pp 60. Edward Arnold, London. James, P. W. & Rose, F. (1976). Survey of lichen habitats in N.W. Sutherland. NCC CST Report, No. 112. NCC, Peterborough. 1977 James, P. W., Hawksworth, D. L. & Rose, F. (1977). Lichen communities in the British Isles: a preliminary conspectus. In: Seaward, M. R. D., ed., Lichen Ecology. Pp 295-413. Academic Press, London. Alvin, K. & Rose, F. (1977). The Observer’s book of lichens. Revised edition. Frederick Warne & Co, London. 1978 Rose, F. (1978). The Observer’s book of British wild flowers. Revised edition. Frederick Warne & Co, London. Bulow-Olsen, A. (1978). Plant Communities (edited and adapted by F. Rose). Penguin Nature Guides. Penguin Books, Harmondsworth. Harding, P. & Rose, F. (1978). Pasture woodlands in lowland Britain and their importance for the conservation of the epiphytes and invertebrates associated with old trees. NCC CST Report No. 211. Institute of Terrestrial Ecology & Kings College, London. Rose, F. (1978). Report on the present state of the Solway mosses and recommendations on their conservation. NCC CST Report No. 224. NCC, Peterborough. 1979 Rose, F., van Haluwyn, C. & Lerond, M. (1979): Itineraire lichenologique en Normandie armoricaine 28 juillet - ler aout 1978. Actes du Museum de Rouen 8: 83-99. Christiansen, M. S., Ramkaer, K., Rose, F. & Sochting, U. (1979): Additions to the Danish lichen flora. Botanisk Tidsskrift 74: 89-115. S27 BIBLIOGRAPHY 1980 Rose, F. (1980). Life style that must survive. Geographical Magazine, June 1980 52 (9): 628-635. 1981 Rose, F. (1981, 2nd edition 1990). The Wild Flower Key. British Isles — NW Europe. pp. 480. Frederick Warne London. 1982 Rose, F. & Roux, C. (1982). Porina stoechadiana Rose et Roux sp. nov.. Bulletin du Musee d Histoire Naturelle de Marseille 42: 69-74. 1983 Rose, F. & Coppins, B. J. (1983). Lichens of Colonsay. Proceedings of the Royal Society of Edinburgh, Section B 83: 403-413. Rose, F. (1983). The vegetative key to wild flowers. Fredrick Warne, London, pp. 48. 1984 Rose, F. (1984). Lichens in Somerset. Nature in Somerset, March 1984: 13-15. Rose, F. (1984). Obituary — Charles Antony Lister. Bulletin Kent Field Club 29: 36. Rose, F. & Wolseley, P. A. (1984). Nettlecombe Park — its history and its epiphytic lichens: an attempt at correlation. Field Studies 6: 117-148. 1985 Coppins, B. J., Rose, F. & Tittensor, R. M. (1985). Lichens from a 16th century Sussex cottage. Lichenologist 17: 297-314. 7 Pentecost, A. & Rose, F. (1985). Changes in the cryptogam flora of the Wealden sandrocks, 1688— 1984. Botanical Journal of the Linnean Society 90: 217-230. 1986 Harding, P. & Rose, F. (1986). Pastures-woodlands in lowland Britain: a review of their importance for wildlife conservation. Institute of Terrestrial Ecology, NERC, Huntingdon. pp. 89. . 1987 Rose, F. (1987). Obituaries: Ronald Archie Boniface (1913-1985). Watsonia 16: 353. Hornby, R. J. & Rose, F. (1987). The use of vascular plants in evaluation of ancient woodlands for nature conservation in southern Britain. NCC report. 1988 Rose, F. (1988). Phytogeographical and ecological aspects of Lobarion communities in Europe. Bot. J. Linn. Soc. 96: 69-79. Rose, F. (1988). Obituaries. Edward Charles Wallace (1909-1986). Watsonia 17: 116-117. Rose, F. (1988). Pitfalls of tetrad recording. B.S.B.I. News 48: 18-19. Rose, F. (1988). Plants to look for in the British Isles some of which might be expected to occur as natives. B.S.B.I. News 49: 11-12. Rose, F. (1988). Gymnadenia conopsea (L.) R.Br. In: Rich, T. C. G. & Rich, M. B. D., eds., Plant Crib. pp. 118-119. BSBI, London. Rose, F. & Walls, R. M. (1988). Oenanthe L., fine-leaved species. In: Rich, T. C. G. & Rich, M. B. D., eds., Plant Crib. pp. 62-63. BSBI, London. Roses ie _ Watson, M. F. & Hawksworth, D. L. (1988). Lichens on elms in the British Isles and the effects of Dutch Elm disease on their status. Lichenologist 20: 327-352. Ferry, B. & Rose, F. (1988). The Dungeness shingle. Ecos 9: 22—24. Rose, F. & Brewis, A. (1988). Cephalanthera rubra (L.) Rich. in Hampshire. Watsonia 17: 176— LWT. Rose, F. (1988). The Great Storm of 16 October 1987 and its effect on the corticolous lichen flora of south-east England. Bulletin British Lichen Society 63: \—2. 1989 Rose, F. (1989). Colour identification guide to the grasses, sedges, rushes and ferns of the British Isles and north-western Europe. Viking, London. pp. 240. Rose, F. (1989). Colour identification guide to the grasses, sedges, rushes and ferns of the British Isles. pp. 240. Viking Penguin, London. Rose, F. (1989). Key to annual Salicornia species of south England and north France (modified from a key by Prof. Gehu (1979) by Dr. F. Rose with some additional material). B.S.B.J. News 53: 12-16. Rose, F. (1989). Obituary. Alice Gertrude Side (1911-1988). Trans. Kent Field Club 11: 47-49. 1990 Rose, F. & Boissiere, J. C. (1990): Catalogue raisonné des lichens du massif de Fontainebleau et des Trois Pignons. Les enseignements de Cette étude. Bulletin de la Société Botanique de France, Lettres Botaniques 137(2/3): 173. BIBLIOGRAPHY S23) Rose, F. (1990). The epiphytic (corticolous and lignicolous) lichen flora of the Forét de Fontainbleau. Bulletin de la Société Botanique de France, Lettres Botaniques 137(2/3): 197— 209. 199] Rose, F., Stern, R. C., Matcham, H. W. & Coppins, B. J. (1991). Atlas of Sussex mosses, liverworts and lichens. Booth Museum, Brighton. Rose, F. (1991). A new subspecies of Gymnadenia conopsea (L.) R. Br. Watsonia 18: 319-320. Rose, F. (1991). Neolithic trees on West Sussex Downs. B.S.B./. News 57: 10-11. 1992 Rose, F. (1992). Temperate forest management; its effects on bryophyte and lichen floras and habitats. In:, Bates, J. & Farmer, A. M. eds., Bryophytes and lichens in a changing environment. pp. 211—233. Clarendon Press, Oxford. Coppins, B. J. & Rose, F. (1992). Maps of Pannariaceae. In: Seaward, M. R. D. (1999). Lichen Atlas of the British Isles. Part 4. British Lichen Society, London. Purvis, O. W. & Rose, F. (1992). Maps of Lecanactis. In: Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. J. & Moore, D. M. (1992). The lichen flora of Great Britain and Ireland. pp. 281—285. Natural History Museum Publications/British Lichen Society, London. Rose, F. & James, P. W. (1992). Pachyphiale. In: Purvis, O. W., Coppins, B. J., Hawksworth, D. L., James, P. J. & Moore, D. M. (1992). The lichen flora of Great Britain and Ireland. pp. 417-418. Natural History Museum Publications/British Lichen Society, London. Rose, F. (1992). Report on the remaining heathlands of West Sussex 1991-92. West Sussex Heathland Forum. West Sussex County Council, Chichester. pp. 48. BoB Rose, F. (1993). Ancient British woodlands and their epiphytes. British Wildlife 5(2): 83-95. Rose, F. (1993). Report on the bryophytes and lichens in chalk grassland in West Sussex. Sussex Downs Conservation Board & West Sussex County Council. pp. 48. McClintock, D. & Rose, F. (1993). Obituaries. Hector Wilks 1919-1992. Transactions Kent Field Club 13: 59-60. Rose, F. (1993). Ancient British woodlands and their epiphytes. British Wildlife 5: 83-93. Rose, F. (1993). Obituary. Thomas Douglas (Dougal) Victor Swinscow (1917-1992). Journal of Bryology 17: 699-700. 1994 Rose, F. (1994). Ancient British woodlands and their epiphytes. British Wildlife 5: 83-93. Rose, F. & James, P. (1994). Lichens rock on at Stonehenge. Plantlife autumn/winter 1994: 8—9. 1995 Rose, F. (1995). Maps of 46 Parmelia species. In: Seaward, M. R. D. (1995). Lichen Atlas of the British Isles. Part 1. British Lichen Society, London. Richardson, S.J., Rose, F., Rich, T.C.G. (1995). The distribution and ecology of Tunbridge Filmy Fern (Hymenophyllum tunbrigense (L.) Smith): Hymenophyllaceae in south-east England in 1994., Plantlife ‘Back from the Brink’ Project Report, 43: pp. 22. Plantlife, London. Rich, T.C.G. & Rose, F. (1995). Tower Mustard (Arabis glabra) in Hampshire in 1995. Plantlife ‘Back from the Brink’ Project Report no. 61. Plantlife, London. Rich, T.C.G., Richardson, S.J. & Rose, F. (1995). Tunbridge Filmy-fern, Hymenophyllum tunbrigense (Hymenophyllaceae: Pteridophyta) in south-east England in 1994/1995. Fern Gazette 15: 51-63. 1996 Brewis, A., Bowman, & Rose, F. (1996). The Flora of Hampshire pp. 386. Harley Books, Colchester. Rose, F. & Davey, S.R. (1996). The three forms of Fragrant Orchid. B.S.B.J. News 72: 71. Rose, F. (1996). The return of the elm. Plantlife spring/summer 1996: 8-9. Rose, F. (1996). The habitats and vegetation of Sussex. pp. 1-27. Booth Museum of Natural History, Brighton. IT) Rose, F. (1997). The current status and conservation issues of cryptogams in the High Weald. In, Jackson, A. & Flannagan, M. eds. 1997. Conservation of cryptogams in the Weald. pp. 19-21. Royal Botanic Gardens, Kew. 1999 Rose, F. & Coppins, B. J. (1999). Maps of Degelia (3 accounts). In: Seaward, M. R. D., ed. (1999). Lichen Atlas of the British Isles. Part 4. British Lichen Society, London. Rose, F. (1999). Indicators of ancient woodland — the use of vascular plants in evaluating ancient woods for nature conservation. British Wildlife 10: 241-251. 524 BIBLIOGRAPHY Rose, F. (1999). The interesting plant life of MOD heathland areas in Hampshire. Sanctuary 28: 20-21. 2000 Rose, F. (2000). Changes in vegetation and flora in the past 1000 years. Transactions of the Suffolk Naturalists’ oe 36: 19-28. Abraham, F. & Rose, F. (2000). Large-leaved limes on the South Downs. British Wildlife 12: 86— 90. 2001 Coppins, B. J., Hawksworth, D. L. & F. Rose (2001). Lichens. Pp 126-147 in Hawksworth, D. L., ed., The changing wildlife of Great Britain and Ireland. The Systematics Association special volume series 62. Taylor & Francis, London. Rose, F. (2001). Finding refuge in God’s acre. Plantlife autumn 2001: 8-9. 2002 Rose, F. (2002). Living in interesting times. Plantlife spring 2002: 10-11. 2006 Rose, F. & O'Reilly, C. (2006, revised and expanded edition). The Wild Flower Key. pp. 576. Frederick Warne, London. UNPUBLISHED REPORTS AND SURVEYS ETC. 1968 Rose, F. & Showell, J. (1968). Windsor Forest, 17/2/68. Manuscript. Report to the Nature Conservancy. Rose, F. (1968). Corticolous and lignicolous lichens of Staverton Park. Manuscript. Pp 4. Nature Conservancy. Rose, F. (1968). Preliminary report on deciduous woodlands and old parklands containing fragments of old deciduous woodland, in Norfolk and Suffolk. I Suffolk. II Norfolk. Pp 20. Manuscript. Report to the Nature Conservancy. Rose, F. (1968). Report on survey carried out in the Forest of Dean, April and May, 1968. Manuscript. Report to the Nature Conservancy. Rose, F. (1968). Survey report on woodland and forest relict areas of the Wealden region. Manuscript. pp. 27. Report to the Nature Conservancy. Rose, F. (1968). Wood south-east of Alcester. Great Wood, Vowchurch. Moccas Park. Banses Wood, south-east of Cockyard. Manuscript. Report to the Nature Conservancy. Rose, F. (1968). Wyre Forest (visited 28th and 29th May, 1968). Manuscript. pp. 21. Report to the Nature Conservancy. 1969 Rose, F. (1969). Kent lichen check-list. Manuscript. Copy with BLS. Rose, F. (1969). Burnham Beeches. Longleat Woods and Park. Cranborne Chase. Savernake Forest. Manuscript. Report to the Nature Conservancy. Rose, F. (1969). Hareshaw Linn, Bellingham. Ingleton Glens, West Riding. Hackfall, West Riding. Manuscript. Report to the Nature Conservancy. Rose, F. (1969). Report to the Nature Conservancy on a visit to Wyre Forest in April 1969 (Supplement to Report of May 1968). Manuscript. Report to the Nature Conservancy. 1970 Rose, F. (1970). Survey report on woodland and forest relict areas of the Wealden region: draft. Nature Conservancy Council, South East Region. pp. 27. Peterken, G. F. & Rose, F. (1970). Felbrigg Wood, Norfolk, pp. 5. IO Rose, F. (1971). Report on surveys of woodland and parkland carried out in Cornwall. 1-4 October 1971. Manuscript. Report to the Nature Conservancy. Rose, F. (1971). A survey of the woodlands of the Lake District, and an assessment of their conservation value based upon structure, age of trees, and lichen and bryophyte epiphyte flora. Manuscript. Report to the Nature Conservancy. Rose, F. (1971). Report on sites visited in East Anglia, June 1970; March 1971; July 1971. Manuscript. Report to the Nature Conservancy. Rose, F. (1971). Report on the cryptogamic vegetation of woodland areas in North Wales Manuscript. Pp 8. Rosey EST) siReporiaon Sotterley Park, East Suffolk. Manuscript. Report to the Nature Conservancy. BIBLIOGRAPHY 525 O72 Rose, F. (1972). Brampton Bryan Park, Herefordshire. Manuscript. pp. 3. Report to the Nature Conservancy. Rose, F. (1972). Downton Castle Park & Gorge, Herefordshire. Manuscript. Report to the Nature Conservancy. Rose, F. (1972). Hull Wood, Glandford. Sheringham Park. Sturston Carr. Hilborough Park. Manuscript. Report to the Nature Conservancy. Rose, F. (1972). Kentchurch Park, Homend Park. Eastnor Park. Holme Lacy Park & Wood, Herefordshire. Manuscript. pp. 4. Report to the Nature Conservancy. Rose, F. (1972). Report on lichenological and bryological survey of the woodlands etc in Morven and the Loch Sunart — Ardnamurchan area. Manuscript. Report to the Nature Conservancy. Rose, F. (1972). Report on Wensleydale sites surveyed Sth and 6th August 1972. Manuscript. Report to the Nature Conservancy. Rose, F. (1972). Survey report on woodland sites in North Yorkshire & Co. Durham. Manuscript. Pp 17. Report to the Nature Conservancy. Rose, F. (1972). Wye Gorge — below Symonds Yat, Herefordshire. Manuscript. pp. 3. Report to the Nature Conservancy. Bowen, H. J. M., Rose, F. & Mason, J. (1972). List of species from Wychwood, Oxon, seen in April NIDA {0s Se 1973 Rose, F. (1973). Report on survey work on cryptogamic vegetation in Essex and Suffolk, 16th—19th March 1973. Manuscript. Report to the Nature Conservancy. Rose, F. (1973). Report on woodland surveys in South Wales, June & September 1972. Manuscript. Pp 9. Report to the Nature Conservancy. Rose, F. (1973). Report on surveys of woodlands in Kintyre — Knapdale and in Sunart with particular reference to their cryptogamic floras. Manuscript. Report to the Nature Conservancy. Rose, F. (1973). Interim report on surveys done in Westmorland and Cumberland August 29th — September Ist, 1973. pp. 6. Manuscript. Report to the Nature Conservancy. 1974 Rose, F. (1974). Survey of woodland and parkland sites in the S.W. Midlands, April 12-18, 1973. Manuscript. pp. 8. Report to the Nature Conservancy. Rose, F. (1974). Report on work done in East Anglia in botanical survey 16th—19th July, 1974. Manuscript. Report to the Nature Conservancy. Rose, F. (1974). Report on survey of woodland and parkland sites in Cumbria, August, 1974. Manuscript. Report to the Nature Conservancy. S75 Rose, F. (1975). Report on woodland sites surveyed in Cumbria September 16th—19th, 1975. Manuscript. Report to the Nature Conservancy. Rose, F. (1975). Report on survey of Tyneham Ranges for bryophytes and lichens, 1&th—20th April, 1975. Manuscript. Report to the Nature Conservancy. Rose, F. (1975). Report on survey of chalk grasslands in the military lands about Tidworth March 13th 1975. Manuscript. Report to the Nature Conservancy. Rose, F. (1975). The lichens and bryophytes of sites in Hereford, Salop, and Staffordshire, based on a survey made from April 21-24, 1975. Manuscript. pp. 10. Report to the Nature Conservancy. Rose, F. (1975). Scales Wood. Manuscript. Report to the Nature Conservancy. Rose, F. (1975). Report on the conservation of epiphytic lichens and bryophytes on the Cranborne Chase Estate, Wiltshire and Dorset. Surveyed August 28 1975.5 pp. Rose, F. (1975). Report on sites surveyed in the (former) Merionethshire, March 27—April 2 1975. Manuscript. Pp 6. 1976 Rose, F. (1976). Report on survey of Cranborne Chase 23 March 1976 and 24 march 1976.. 5 pp. Rose, F. & James, P. W. (1976). A survey of the lichens of Galloway and western Dumfries-shire. Nature Conservancy Council, Scotland, South West Region. pp. 25. Rose, F. (1976). Report on survey of cryptogam flora (bryophytes & lichens) of the Longleat Estate surveyed August 1969, November 1973 & 27 November 1974. Manuscript. pp. 4 and pp. 2. Report to the Nature Conservancy. O77 Rose, F. (1977). Chequers Park. pp 1. Rose, F. (1977). Stonor Park. pp.1. 526 BIBLIOGRAPHY 1981 Rose, F. & James, P. W. (1981). Report on the lichen flora and vegetation of lowland areas of the Cowal peninsula, 21—30 June 1977. Nature Conservancy Council. Scotland, South West Region. pp. 19. 1983 Rose, F. (1983). Report on the populations of Ophrys sphegodes in South East England in May 1983. Nature Conservancy Council. England. South East Region. pp. 21. 1986 Hornby, R. J. & Rose, F. (1986). The use of vascular plants in evaluation of ancient woodlands for nature conservation in southern England. Nature Conservancy Council, Peterborough. pp. 12. 1988 Rose, F. (1988). Report on lichen surveys of sites in West Sussex to assess the effects of the Hurricane of October 16, 1987. Nature Conservancy Council. England. South East Region. pp. 10. Rose, F. & Yates, E. M. (1988). The vegetation of Ebernoe Common, Sussex: an historical and botanical survey. King’s College London. Rogate Field Centre. pp. 20. Rose, F. (1989). A survey of Tickfold Ghyll — West Sussex. Nature Conservancy Council. 1989 Rose, F. (1989). Report on lichen and bryophyte survey of Ashburnham Park SSSI, East Sussex, March-— —April 1989. No place of publication given. 199 Rose, F. (1991). A survey of the bryophytes in Fairlight Glen, East Sussex. English Nature, South East Region. pp. 8. Rose, F. (1991). A survey of lichens and bryophytes at Eridge Park SSSI. English Nature, South East Region. pp. 10. 1992 Rose, F. (1992). Bryophytes and pteridophytes on Wealden Sandrocks, Philpots Estate, Chiddingly Wood. English Nature, South East Region. pp. 6. 1994 Rose, F. (1994). Report on surveys of woodlands in Cornwall with particular attention to their epiphytic lichen floras during May and September/October, 1992 and also over period 1972— 199]. English Nature, South West Region. NCC, Peterborough. 1996 Cox, J. & Rose, F. (1996). Species Recovery Project: summer lady’s-tresses. English Nature Species Recovery Programme & Hampshire Wildlife Trust. pp. 39. 1997 Rose, F. (1997). Report on the present and past botanical interest of the Open Pits on Dungeness Beach, with suggestions for future management. English Nature, Kent Team. pp. 16. Rose, F. & Patmore, J. (1997). Weald gill woodlands. English Nature, Sussex and Surrey Local Team. 1998 Rose, F. & Davy, S. (1998). 4 lichenological survey of Ethy Park. National Trust. pp. 49. INSTRUCTIONS TO CONTRIBUTORS Scope: Authors are invited to submit Papers and Notes concerning British and Irish vascular plants, their taxonomy, biosystematics, ecology, distribution and conservation, as well as topics of a more general or historical nature. Authors should consult the Hon. Receiving Editor for advice on suitability or any other matter relating to submission of manuscripts. Each paper is handled by one of the Editorial Panel who will usually seek comments from an appropriate referee before acceptance. Papers and Notes must be submitted in duplicate, typewritten on one side of the paper, with wide margins and double-spaced throughout. Pages should be numbered. 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Sanford, c/o The Museum, High Street, Ipswich, Suffolk, IP] 3QH. Books for Review: John Atkins, Summerfield House, High Street, Brough, Kirkby Stephen, Cumbria CA17 4AX. Plant Records: the appropriate vice-county recorder, who should then send them to Mike Porter, 5 West Avenue, Wigton, Cumbria CA7 9LG. Obituaries: Mrs M. Briggs, 9 Arun Prospect, Pulborough, West Sussex, RH20 IAL. Back issues of Watsonia Available from BSBI Hon. Membership Secretary, Gwynn Ellis, 41 Marlborough Road, Roath, Cardiff, CF2 5BU. E-mail: rgellis@ntlworld.com Tel & Fax 029 2049 6042 warsonia MUTI Contents PRICE, D. T. & RICH, T. C. G. One-way introgressive hybridisation between Sorbus aria and S. torminalis (Rosaceae) in southern Britain Copp, R. B. & RICH, T. C. G. Variation in ae Biieiracee (Rubiaceae) i in the British Isles .. SLADE, [Ky é& RICH, To €:°G: poles agi in British one sect. Alpina (Asteraceae) RicH, T..C. 'G., EDWARDS, B. & PEARMAN, D. A. eee wom (Asteraceae), a new endemic hawkweed from the Isle of Portland, wate. related to Hieracium leyanum RicH, T. €.G. & SMITs,P. At Caan of Britain’s s biodiversity Hanae cacuminum, Summit Hawkweed (Asteraceae) . KITCHENER, G. D. Rumex patientia L. x R. conglomerat Muay a newly described dock hybrid RUMSEY, F. J. A reconsideration of Dealieaie maritima are (Orobanchacese) and related taxa in southern England and the Channel Islands RUMSEY, F. J. An overlooked boreal clubmoss Lycopodium = as ex Hartm.) Zinserl. ex Kusen. (Lycopodiaceae) in Britain MARGETTS, L. J. A new species of Taraxacum Wigg. pacar gas aie west England and Ireland a tates Se Wesel eg een ana ae NOTES Allen, D. E. A fourth list of British species of Rubus L. casa in north-west France}: aaa ie sh WOR ae an Bec is, tae 485-486 Nelson, E. CC. Brica x ee Druce cE. tetralix L. x vagans ie anote ontypification .... 487-488 Rumsey, F. J. An early specimen of Cystopers diaphana (Bory) Blasdell Suppo its native status re 488-490 Rich, 1. C. G., Sawtschuk, 1 c ee iP. The British mdse Hevea cambricogothicum (Asteraceae) is probably extinct . wae” deat ae 490-491 PLANT RECORDS) is.) ces: ceay ea See ee 493-510 OBITUARIES _... Bs: se ee ie ane mee , es) ee siya EE 511-526 ISSN 0043-1532 Typeset by D. K. & M.N. SANFORD Printed in Great Britain by HENRY LING LIMITED, THE DORSET PRESS, DORCHESTER, DORSET DT1 1HD Ary Be SMITHSONIAN INSTITUTION LIBRARIES Ml WNL 3 9088 01359 4924 eae 5 Ag Byes ses tz -, ers oe arate, Me EK nag set OT EST bed St oe ire Sawa ee PqE SE SS oT See ere ere 2 St? = ES ers: eS ite See nity a ’